from Miocene Mexican amber

Palaeontographica, Abt. A: Palaeozoology – Stratigraphy Vol. 303, Issues 1–3: 1–21 Stuttgart, September 2014

Article

Systematic and palaeoecology of Psychodidae (Insecta: Diptera) from Miocene Mexican amber With 2 plates, 7 text-figures and 1 table

Frauke Stebner and Mónica M. Solórzano Kraemer Abstract Psychodidae is a family of nematoceran flies distributed worldwide with about 3000 extant and more than 90 fossil species from diverse Tertiary and Cretaceous ambers. However, the fossil and also recent psychodid fauna from Mexico is still poorly investigated although Mexican amber is one of the most important amber deposits of the Tertiary and the region is considered to belong to a megadiversity area today. In the present work 40 fossil Psychodidae in Mexican amber from Simojovel are studied with the focus on systematic descriptions and palaeoecological significance. The composition of the fossil psychodid fauna with primarily subaquatic and terrestrial genera but only a few aquatic representatives indicates a tropical forest with damp ground containing masses of decomposing organic matter. Six new species and one new genus are described: Trichomyia richardi n. sp., Trichomyia axeli n. sp., Psychoda rusti n. sp., Psychoda tzotzili n. sp., Philosepedon apozonallii n. sp. and Matuna bechlyi n. gen. et sp. Key words: Psychodidae, Mexican amber, Miocene, systematics, palaeoecology

Zusammenfassung Die Psychodiden sind eine weltweit verbreitete Familie der Dipteren mit mehr als 3000 rezenten und über 90 fossilen Arten aus tertiären und kretazischen Bernsteinen. Obwohl der miozäne Mexikanische Bernstein zu einem der wichtigsten Bernstein Vorkommen des Tertiärs zählt und die Region heute ein Megadiversitäts-Gebiet darstellt, ist die fossile sowie die rezente Psychodiden-Fauna nur wenig untersucht. In der vorliegenden Arbeit werden 40 fossile Psychodiden aus dem Mexikanischen Bernstein systematisch und paläoökologisch untersucht. Die Zusammensetzung der fossilen Psychodiden-Fauna mit vornehmlich subaquatischen und terrestrischen, aber nur wenigen aquatischen Vertretern, deutet auf einen tropischen Wald hin, der durch feuchte Böden mit einem hohen Totholzanteil und verrottendem Pflanzenmaterial charakterisiert war. Sechs neue Arten und eine neue Gattung werden beschrieben: Trichomyia richardi n. sp., Trichomyia axeli n. sp., Psychoda rusti n. sp., Psychoda tzotzili n. sp., Philosepedon apozonallii n. sp. und Matuna bechlyi n. gen. et sp. Schlüsselwörter: Psychodidae, Mexikanischer Bernstein, Miozän, Systematik, Paläoökologie

Table of contents 1. Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2. Material and Methods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3. Systematic descriptions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Subfamily Trichomyiinae . . . . . . . . . . . . . . . . . . . . . . . . . . . . Genus Trichomiyia Haliday in Curtis, 1839 . . . . . . . Subfamily Psychodinae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Genus Psychoda Latreille, 1796 . . . . . . . . . . . . . . . . .

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Genus Philosepedon Eaton, 1904 . . . . . . . . . . . . . . . . . . Matuna n. gen. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4. Discussion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5. Acknowledgements . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6. References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Explanation of plates . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

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Authors’ addresses: Frauke Stebner (corresponding author), Steinmann-Institute, University of Bonn, Nussallee 8, D-53115 Bonn, Germany, e-mail: [email protected], phone: +47/228/733336. Mónica M. Solórzano Kraemer, Steinmann-Institute, University of Bonn, Nussallee 8, D-53115 Bonn, Germany and Senckenberg Research Institute and Natural History Museum Frankfurt, Senckenberganlage 25, 60325 Frankfurt, Germany © 2014 E. Schweizerbart’sche Verlagsbuchhandlung, Stuttgart, Germany

www.schweizerbart.de

0375-0442/14/0303-0046

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$ 5.25

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Frauke Stebner and Mónica M. Solórzano Kraemer

1. Introduction Psychodidae is a large family of nematoceran flies with currently 3026 described species (Pape et al. 2011) in six subfamilies, including Psychodinae, Trichomyiinae, Bruchomyiinae, Sycoracinae, Horaiellinae, and Phlebotominae, although the latter one has been given family rank by some authors (Rohdendorf 1964, Abonnenc & Leger 1976, Williams 1993). Adults commonly occur in moist and wooded areas and the larvae inhabit various aquatic to terrestrial environments. The feeding habit of adults varies greatly. While females of Phlebotominae and Sycoracinae are blood feeding, adults of the remaining groups probably do not feed at all. Today Psychodidae are distributed worldwide except Antarctica. In Central America they are represented by about 274 species (Wagner & IbáñezBernal 2009). With 65 recorded extant species the psychodid fauna of Mexico is still poorly investigated but probably only 20% of all species are already known (Ibáñez-Bernal 2000). The fossil record of Psychodidae contains more than 90 species (Evenhuis 1994, EDNA Fossil Insect Database, accessed on August 2013: http:// edna.palass-hosting.org). The oldest record is from the Lower Jurassic of northeastern Germany (Ansorge 1994) and some of the oldest amber Psychodidae are from the Lower Cretaceous Lebanese amber (Azar et al. 1999, Azar et al. 2003a, Azar et al. 2003b) and the Lower Cenomanian French amber (Azar et al. 2007, Lak et al. 2008). To date only 14 fossil species have been described from Mexican amber (Quate 1961, Quate 1963, Ibáñez-Bernal et al. 2013, Coty et al. 2013). In the present study 40 psychodids in Mexican amber have been studied and compared to fossil and extant species with the main focus on systematic descriptions and their palaeobiology and -ecology. The 39 fossils investigated belong to three subfamilies: Trichomyiinae (18), Psychodinae (20) and Phlebotominae (1). The classification of recent genera is sometimes problematic and controversial within the Psychodinae and Trichomyiinae and due to the state of preservation the assignment of fossils is even more problematic. However, within this work 30 of the 39 fossils could be assigned to five recent genera and one new genus, Trichomyia (18), Psychoda (5), Threticus (3), Alepia (1), Philosepedon (1), and Matuna n. gen. (2), and six new species are described: Trichomyia richardi n. sp., Trichomyia axeli n.

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sp., Psychoda rusti n. sp., Psychoda tzotzili n. sp., Philosepedon apozonallii n. sp. and Matuna bechlyi n. gen. et sp. The only phlebotomine specimen has been described prior to this study as Pintomyia bolontikui (Ibáñez-Bernal et al. 2013). Because modern representatives are restricted to certain ecological conditions the investigation of fossils allows at least partly reconstruction of the former ecosystem of the extinct amber forest. For the Mexican amber a tropical dry lowland forest with a mangrove region has been reconstructed (Solórzano Kraemer 2007). In this context Mexican amber is of particular interest because of the existence of an extant forest located in the south-eastern part of Mexico at the Pacific coast in the state of Chiapas with a floral composition quite similar to the past amber forest, as shown by fossil pollen analysis of the amber bearing sediments (Langenheim 1967, MartínezHernández 1992, Graham 1999). Therefore the analysis of the fossil fauna from amber in comparison to the fauna from the extant amber forest provides opportunities for future ecological studies. Finally, the study of fossil insects from Mexican amber is a contribution to the knowledge of biodiversity of a highly diverse but still poorly investigated region.

2. Material and Methods The present work is based on 40 individuals of Psychodidae from Miocene Mexican amber (see Table 1). The amber samples are loaned from collections of the University of California, Museum of Paleontology, Berkeley, California, USA (hereafter UCMP), and the Stuttgart State Museum of Natural History Schloss Rosenstein, Stuttgart, Germany (hereafter SMNS). Inventory numbers and specimen numbers are listed in Table 1. The morphological terminology follows Quate & Vockeroth (1981). The following abbreviations are used in the systematic descriptions and text-figures: aed: aedeagus; asc: ascoid; cer: cerci; CuA1/ CuA2: branches of cubitus; d proj: dorsal projection of gonopod; goncx: gonocoxite; gonst: gonostylus; M1–M3: branches of media; M fk: medial fork; par: paramere; ped: pedicel; R1–R5: branches of radius; R fk: radial fork; r-m: radial-medial crossvein; Sc: subcosta; scp: scape; surst: surstylus; tenac: tenaculum; v proj: ventral projection of gonopod.

Systematic and palaeoecology of Psychodidae

Table 1. List of Psychodidae in Mexican amber from the collection of the UCMP and the SMNS.

Collection UCMP UCMP

UCMP Specimen Number 10375 10376

Inventory Number B-6899-41

Subfamily Psychodinae

B-6899-42

Psychodinae ♀

UCMP

10377

B-6899-61

Psychodinae

Threticus sp. ♂

UCMP

10378

B-7044-27

Trichomyiinae

Trichomyia sp. ♀

UCMP

10379

B-7044-29

Psychodinae

Threticus sp. ♂

UCMP UCMP UCMP

10380 10381 10382

B-7045-38 B-7047-32

Psychodinae Trichomyiinae

Psychoda Trichomyia

B-7048-38

Trichomyiinae

Trichomyia sp. ♂

UCMP

10383

B-7048-39

Trichomyiinae

Trichomyia sp. ♀

Genus Matuna n. gen.

Species Matuna bechlyi n. sp. ♂

Psychoda tzotzili n. sp. ♂ Trichomyia richardi n. sp. ♂

UCMP

10384

B-7053-33

Trichomyiinae

Trichomyia sp. ♀

UCMP

10385

B-8411-57

Psychodinae

Psychoda sp. ♂

UCMP

10386

B-8411-61

Trichomyiinae

Trichomyia sp. ♂

UCMP

10387

B-8413-146

Psychodinae ♂

UCMP

10388

B-8413-149

Psychodinae

UCMP UCMP UCMP

10389 10390 10391

B-8416-6

Trichomyiinae

Trichomyia

Trichomyia axeli n. sp. ♂

B-8419-27

Psychodinae

Philosepedon

Philosepedon apozonallii n. sp. ♂

B-8419-28

Trichomyiinae

Trichomyia sp. ♀

UCMP

10392

B-8419-29

Trichomyiinae

Trichomyia sp. ♀

B-8421-10

Trichomyiinae

Trichomyia sp. ♀

Threticus sp. ♂

UCMP

10393

UCMP

10394

B-8421-11

Trichomyiinae

Trichomyia sp. ♀

UCMP UCMP

10395 10396

B-8425-92

Psychodinae

Psychoda

B-8425-93

Trichomyiinae

Trichomyia sp. ♂

UCMP

10397

B-8425-94

Trichomyiinae

Trichomyia sp. ♀

UCMP

10398

D-698-24

Trichomyiinae

Trichomyia sp. ♀

Mx 118

Psychodinae ♂

SMNS

Psychoda rusti n. sp. ♂

Psychodinae ♂ SMNS

Mx 216

Psychodinae

Psychoda sp. ♂

SMNS

Mx 217

Trichomyiinae

Trichomyia sp. ♀

SMNS

Mx 220

Psychodinae

Psychoda sp. ♀

SMNS

Mx 226

Trichomyiinae

Trichomyia sp. ♀

SMNS

Mx 229

Trichomyiinae

Trichomyia sp.

SMNS

Mx 235

Psychodinae

Matuna sp. ♂

Psychodinae ♀ SMNS

Mx 276

Psychodinae Psychodinae

SMNS

Mx 278

Trichomyiinae

SMNS

Mx 304

Psychodinae ♀

SMNS

Mx 346

Psychodinae

Alepia sp. ♀

SMNS

Mx 351.3

Phlebotominae

Pintomyia

Trichomyia sp.

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Pintomyia bolontikui ♂

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For the taxonomic identification and investigation a Leica MZ 125 was used. Drawings were rendered with the aid of a drawing tube and measurements are given in millimeters. Photographs of the amber inclusion were made with a Leica MZ 16 Stereomicroscope with a JVC ky-F70B Digital Camera. Compound photographs merging different focus levels to a single image were performed by using Discus software equipped with stacking function. Since even the association of recent males and females is often problematic, descriptions of new species in the present work are based on males only.

3. Systematic descriptions Subfamily Trichomyiinae Genus Trichomyia Haliday in Curtis, 1839 Trichomyia richardi n. sp. (Plate 2, Figs A, E; Text-fig. 1 A–C) Type material: Holotype is a male from the collection of the UCMP with inventory number B-7047-32. Type locality: Early Middle Miocene amber from Simojovel de Allende, Chiapas, Mexico.

Text-fig. 1. Trichomyia richardi n. sp. (A) wing. (B) genitalia, lateral view. (C) antenna: pedicel and flagellomeres 1–3.

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Etymology: The specific epithet is dedicated to Richard Stebner (Bonn).

Diagnosis male: Genitalia with gonopod comprising two lateral projections, dorsal projection digitiform, longer than ventral one, ventral projection bifurcate; aedeagus reaching to middle of dorsal projection, with two curved filament-like structures arising apically and with one spine-like structure on each side. Description male: Head: Flattened, distinctly wider than long; with frontal and vertical setae curving forward. Eyes projecting over lower part of head, rather oval in shape, with postocular bristles on upper margin. Both antennae broken, each with 11 flagellomeres. Scape cylindrical; pedicel spherical with three sensilla originating from lower basal part of pedicel. Flagellomeres (Text-fig. 1 C) asymmetrical pyriform and recessed, first flagellomere more nearly cylindrical. Each flagellomere with a pair of curved simple ascoids; ascoids distinctly longer than flagellomeres bearing them, reaching distal third of following flagellomere. Basal two thirds of each flagellomere covered with fine hair. Relative length of antennomeres 1–13: 10 – 12.5 – 16.5 – 15 – 14 – 14.5 – 14 – 12.5 – 11.5 – 11.5 – 11.5 – 11 – 9. Maxillary palpus three-segmented, no depression visible. Wing (Text-fig. 1 A): Broad with rounded apex; vestiture confined to veins. Wing length: 1.2 mm; greatest width: 0.5 mm. Sc ends into costa and at its tip is a crossvein towards R1. Medial fork at centre of wing; radial fork slightly distad of medial fork. Medial fork at level of tip of CuA2; radial fork between level of tip of CuA1 and CuA2. R5 ends at rounded wing apex. Basal part of CuA2 widened. Crossvein r-m present but faint. Abdomen: Tergum 1 and sternum 1 reduced to a narrow ring; tergum 2 half as wide as following one; tergum 7 with two lateral lobes and a median groove. Genitalia (Text-fig. 1 B): Posterior edge of tergum 9 with two triangular protuberances flanking a Vshaped median groove. Gonopod with two lateral projections; dorsal projection (gonocoxite?) digitiform, longer than ventral one, with long hairs on entire upper margin; ventral projection (gonostylus?) bifurcate. Dorsal part of ventral projection curving inwards, inflated at centre and with an acute tapered apex, ventral part digitiform, with an angular apex and three long setae. Aedeagus reaching to middle of dorsal projection, situated dorsally, with two curved filament-like structures on apex and one spine-like structure on each side (parameres?), beneath those two

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smaller spine-like structures appear. Cerci broadened from basal part to apex in lateral view, apex angular. Remarks: Trichomyiinae are represented by two fossil Cenozoic genera, Eatonisca Meunier, 1905, and Eotrichomyia Nel et al., 2002, and only one recent genus, Trichomyia Haliday in Curtis, 1839, which contains a large number of species with a wide range of morphological disparity and certainly needs to be revised. Bravo (1999) erected the subgenus Septemtrichomyia for neotropical species based on the shape of tergum seven with expanded lateral lobes and various numbers of bristles. Due to the shape of the seventh tergite Trichomyia richardi n. sp. resembles species of Septemtrichomyia but T. richardi n. sp. lacks the bristles on tergum seven and is therefore not included in this subgenus. Based on the unique and complicated shape of the male gonopod the species probably belongs to a new genus of Trichomyiinae but is described here as a new species within the genus Trichomyia until there is a stable and revised classification of Trichomyiinae. Trichomyia axeli n. sp. (Plate 2, Figs D, F; Text-fig. 2 A–C) Type material: Holotype is a male from the collection of the UCMP with inventory number B-8416-6. Type locality: Early Middle Miocene amber from Simojovel de Allende, Chiapas, Mexico. Etymology: The specific epithet is dedicated to Axel Olearius (Frankfurt).

Diagnosis male: Genitalia with gonopod comprising two lateral projections, dorsal projection digitiform and longer than ventral one, ventral projection bifurcate, dorsal part of ventral projection curving inwards, ventral part straight, with three long apical setae. Cerci broadened from basal part to angular apex. Description male: Head: Slightly wider than long, with dense frontal and vertical setae curving downwards. Eyes oval without bridge, with postocular setae. Antennae probably broken; left antenna with 13 antennomeres; right antenna with 14 antennomeres. Scape approximately cylindrical; pedicel slightly bulbous. Flagellomeres (Text-fig. 2 C) pyriform, first flagellomere almost symmetrical, the following ones eccentric. Basal two thirds of each flagellomere covered with hairs. Ascoids stout and curved, longer than flagellomeres bearing them, originating rather basal on flagellomeres. Relative length of antennomeres 1–13 (left antenna): 10 – 12 – 17 – 16 – 15 – 16 – 16 – 14 – 14 – 14 – 17 – 16 – 16. Maxillary palpus three-seg-

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mented, with basal segment distinctly longer than following ones, no depression visible. Relative length of segments: 10 – 7.5 – 6.5. Thorax: With postpronotal setae. Wing (Text-fig. 2 A): With vestiture confined to veins. Length: 1.1 mm, width: 0.4 mm. Sc ends into costa and at its tip is a crossvein towards R1. Medial fork at centre of wing; radial fork distad of medial fork. Radial fork between tip of M3 and CuA1; medial fork between tip of CuA1 and CuA2. Crossvein r-m present. R5 ends approximately at rounded apex of wing. Abdomen: With sparse vestiture. Sterna one and six reduced. Tergum 1 reduced to a narrow ring; tergum 2 approximately half as broad as following one. Tergum 7 with two lateral lobes posteriorly, giving it an U-shaped appearance. Tergum 8 retracted or reduced to a narrow clasp.

Genitalia (Text-fig. 2 B): Posterior edge of tergum 9 with a V-shaped median groove. Gonopod comprising two lateral projections; dorsal projection (gonocxite?) digitiform, longer than ventral one and with long setae on upper surface, curving inwards; ventral projection (gonostylus?) narrower and shorter than dorsal one, bifurcate. Dorsal part of ventral projection curving inwards, inflated at centre; ventral part straight, with three long apical setae. From between the dorsal projection two spines appear (parameres?), narrowing from base to acute apex, curving upwards. Cerci broadened from basal part to apex, apex angular and with numerous short hairs around margin. Aedeagus not clearly visible but apparently not situated between the dorsal projection of gonopod but rather ventral. Remarks: Trichomyia axeli n. sp. shares with Trichomyia richardi n. sp. the following combination of

Text-fig. 2. Trichomyia axeli n. sp. (A) wing. (B) genitalia, lateral view. (C) antenna: scape, pedicel and flagellomeres 1–2.

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characters of the male genitalia: gonopod comprising two lateral projections, dorsal projection digitiform, longer than ventral one, ventral projection bifurcate. The two species differ in the shape and position of the aedeagus which arises from between the dorsal projection of the gonopod in T. richardi n. sp. and bears two curved apical filament-like structures while the aedeagus is situated rather ventrally in T. axeli n. sp. The two species share the shape of tergum 7 with species of the subgenus Septemtrichomyia but differ in lacking the bristles on tergum 7. The two species are closely related and probably belong to a new genus of Trichomyiinae based on the distinct shape of the gonopods,

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but they are described as two new species within the genus Trichomyia until a revision and stable classification of Trichomyiinae is available. Subfamily Psychodinae Genus Psychoda Latreille, 1796 Psychoda rusti n. sp. (Plate 1, Fig. B; Text-fig. 3 A–C) Type material: Holotype is a male from the collection of the UCMP with inventory number B-8425-92. Type locality: Early Middle Miocene amber from Simojovel de Allende, Chiapas, Mexico.

Text-fig. 3. Psychoda rusti n. sp. (A) wing. (B) genitalia, lateral view. (C) antenna: flagellomeres 11–14.

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Etymology: The specific epithet is dedicated to Prof. Dr. Jes Rust (University of Bonn).

Diagnosis male: Flagellomeres 11–14 separated; genitalia with gonostyli short, slightly bent, tapering from base to tip, surstyli short, each with one long and slender tenaculum with acute apex. Description male: Head: Wider than long; eyes reniform, with four medial facet rows; distance between eyes increasing from one facet diameter in upper row to four facet diameters in lower row. No interocular suture. Frons without setae. Antenna with 16 antennomeres; scape cylindrical, approximately twice as long as wide and more than two times longer than pedicel; pedicel spherical. Flagellomeres nodiform with well-developed nodes little longer than internodes; nodes progressively decreasing in size and internodes increasing in length. Flagellomeres with a pair of Y-shaped ascoids, originating from distal part of bulbous base, the two posterior branches extend to base of following node. Flagellomeres 11–14 distinctly separated from each other but with very short internodes; terminal three flagellomeres (12–14) without ascoids, reduced in size, terminal flagellomere dropshaped, somewhat smaller than previous two (Textfig. 3 C). Maxillary palpus not clearly visible, apparently four-segmented, with segments apparently subequal in size. Labellum flattened. Wing (Text-fig. 3 A): Folded so that basal part of the wing with Sc and vein CuA2 is not visible. Length: 1.2 mm, approximately twice as long as wide. No vestiture visible. Forks complete, before centre of wing, radial fork slightly distad of medial fork. R5 ending in apex of wing. Thorax and abdomen: Vestiture lost. Genitalia (Text-fig. 3 B): Gonocoxites cylindrical; gonostyli short, approximately as long as gonocoxites, slightly bent, basal two third tapering evenly, then narrowing abruptly and ending in acute apex. Tergum 9 rectangular, wider than long. Surstylus short, little longer than gonostylus and 1.2 times longer than tergum 9; basal half stout; distal half cylindrical in lateral view, with a curved acute apex, bearing one long and slender tenaculum with acute apex. Aedeagus asymmetrical with tube shaped distiphallus. Remarks: The described specimen shares with modern representatives of Psychoda (sensu Quate 1996) the following combination of characters and can therefore clearly be assigned to this genus: eye-bridge with four rows of facets; interocular suture absent; antenna with 12-14 flagellomeres, those beyond 11 re-

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duced in size, ascoids Y-shaped; labellum flattened; male genitalia with aedeagus asymmetrical, surstylus with single tenaculum. Psychoda rusti n. sp. resembles the two extant species Psychoda cinerea Banks, 1894, and Psychoda tothastica Quate, 1955 in the following characters: antenna with 14 flagellomeres, last three flagellomeres subequal in size and separated from each other; surstylus short, approximately as long as gonostylus. There are, however, differences in the gonostyli, which are shorter in P. tothastica, the shape of the tenaculum which has an acute apex in P. rusti n. sp. and the distance between the eyes, which is less in P. cinerea and P. tothastica (one to two facet diameters in P. cinerea and one facet diameter in P. tothastica). The shape of the gonopod of P. rusti n. sp. is very similar to P. cinerea but there are differences in the shape of the gonostylus which is apically more bent and tapers evenly in P. cinerea. Psychoda tzotzili n. sp. (Plate 1, Figs A, D; Text-fig. 4 A–D) Type material: Holotype is a male from the collection of the UCMP with inventory number B-7045-38. Type locality: Early Middle Miocene amber from Simojovel de Allende, Chiapas, Mexico. Etymology: The specific epithet refers to the Tzotzil, the indigenous group of inhabitants of the central Chiapas highlands in southern Mexico.

Diagnosis male: Flagellomeres 11–13 without internodes but separated; genitalia with gonostylus curved inwards, with one long prebasal seta, surstylus with basal protuberance and tube shaped distal part, with a single club shaped tenaculum, surstyli one and a half times longer than tergum 9 and two times longer than gonocoxite. Description male: Head: Eyes reniform, with four medial facet rows; distance between eyes not clearly visible, but apparently increasing from upper to lower margin. Frons without setae; vertical setae short and curved backwards. Right antenna broken, with 11 antennomeres; left antenna with 15 antennomeres, about half as long as wing; scape cylindrical, one and a half times longer than wide and longer than bulbous pedicel (Text-fig. 4 C). Scape and pedicel covered with fine hair; pedicel with two setae on upper surface. Flagellomeres nodiform with well-developed nodes; node of first flagellomere slightly longer than internode; nodes progressively decreasing in size, internodes increasing in length. Flagellomeres 1–11


each with a pair of Y-shaped ascoids, the two anterior branches reaching the level of the base of the following flagellomere; flagellomeres 11-13 without internodes but separated from each other by a suture; terminal two flagellomeres reduced and subequal in size, half the size of the previous one (Text-fig. 4 D). Maxillary palpus four-segmented, about one third as long as antenna, segments equal in length. Labellum not visible. Wing (Text-fig. 4 A): Length 0.9 mm, two and a half times longer than wide, sparse vestiture confined to veins, but probably a lot of vestiture lost because of accumulation of loose hair around wing. Wing venation very faint, especially at basal part and around centre of wing, so that Sc and medial fork are not visible. Radial fork at centre of wing, medial fork must be at

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least at same level or basad of radial fork. R5 ending in apex of wing. Genitalia (Text-fig. 4 B): Gonocoxites with ventral protuberance, straight at inner margin. Apex of gonostyli curved inwards, gonostyli crossing at midline, ending in acute tip, with one long prebasal seta. Surstyli with basal swelling, distal two thirds approximately cylindrical, bearing a single club shaped tenaculum; surstyli one and a half times longer than tergum 9 and two times longer than gonocoxite. Aedeagus not clearly visible, distiphallus tube shaped with angular apex. Two small tube shaped structures (parameres?) with rounded apices appear from beneath the aedeagus.

Text-fig. 4. Psychoda tzotzili n. sp. (A) wing. (B) genitalia, dorsal view. (C) antenna: scape, pedicel and flagellomeres 1–2. (D) antenna: flagellomeres 10–13.

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Remarks: The specimen can be assigned to the genus Psychoda (sensu Quate 1996) by possessing the following combination of characters: antenna with 12– 14 flagellomeres, those beyond 11 reduced; ascoids Y-shaped; eye-bridge with four rows of facets; surstylus with single tenaculum. Psychoda tzotzili n. sp. resembles the extant species Psychoda phalaenoides Linnaeus, 1758, Psychoda uniformata Haseman, 1907, and Psychoda zetoscota Quate, 1959 in the shape of the terminal flagellomeres with flagellomeres 11–13 without internodes but separated from each other by a suture. The males of P. uniformata and P. zetoscota are unknown and the description is based on females only. However, the wing

venation of P. zetoscota differs from P. tzotzili n. sp. in lacking the bases of R3 and M2. The main difference between P. tzotzili n. sp. and P. phalaenoides is the shape of the gonocoxites and surstyli, which are narrower in P. phalaenoides so that the above specimen is described as a new species. Genus Philosepedon Eaton, 1904 Philosepedon apozonallii n. sp. (Plate 1, Fig. E; Text-fig. 5 A–C) Type material: Holotype is a male from the collection of the UCMP with inventory number B-8419-27. Type locality: Early Middle Miocene amber from Simojovel de Allende, Chiapas, Mexico.

Text-fig. 5. Philosepedon apozonallii n. sp. (A) wing. (B) genitalia, lateral view. (C) antenna: scape, pedicel and flagellomeres 1–2.

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Etymology: The specific epithet refers to the Mexican word apozonalli (Nahuatl language) which means amber.

Diagnosis male: Genitalia with gonocoxites flattened; gonostyli bent, tapering from base to acute tip; surstylus ending in rounded apex with one terminal and one subterminal club-shaped tenaculum. Description male: Head: With short vertical setae and a row of postocular setae; eyes reniform with four median facet rows, distance between eyes equals one facet diameter. Both antennae broken, right antenna with nine, left antenna with 10 remaining antennomeres; scape cylindrical and 1.3 times longer than spherical pedicel. Flagellomeres (Text-fig. 5 C) nodiform with well-developed nodes, node one 1.25 x longer than internode, nodes progressively decreasing in size, internodes progressively increasing in length. Each node bearing one pair of Y-shaped ascoids with two anterior branches and one posterior branch. Maxillary palpus four-segmented, about 0.4 mm in length, ratio of the segments: 10 – 13.3 – 13.3 – 15.8. Labellum bulbous. Wing (Text-fig. 5 A): With vestiture confined to veins. Length: 1.4 mm, greatest width 0.6 mm. Forks complete, medial fork before centre of wing, radial fork distad of medial fork, nearly at centre of wing. R5 ending in acute wing apex. Crossvein r–m present. Abdomen: Terga densely covered with hairs. Genitalia (Text-fig. 5 B): Gonocoxites conical and flattened, covered with hairs; gonostyli slightly bent, tapering evenly from base to acute tip. Surstylus hirsute, without distinct protuberance, ending in rounded apex with one terminal and one subterminal calvate tenaculum. Surstyli about two times longer than tergum 9. Aedeagus symmetrical, distiphallus tube shaped, tapering distally to rounded apex. Remarks: The tribal classification and generic definition of Philosepedon is unclear. Philosepedon helicis for example has been considered to belong to the genus Eurygarka by some authors based on the foliaceous ascoids (see discussion in Ibáñez-Bernal & Cáceres 2005 and Curler & Moulton 2008). According to Quate (1958, 1996) the genus can be recognized by the following characters: eye bridge usually with 4 rows of facets; interocular suture present; labellum bulbous, with long spines but without teeth; usually 14 flagellomeres, those beyond 11 reduced and usually subequal and separated; ascoids Y-shaped. Male genitalia with symmetrical aedeagus; surstylus usually with one tenaculum, but may have up to seven. However, most species of Philosepedon have two te-

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nacula or two groups of tenacula (Vaillant 1973, Ibáñez-Bernal 2004a). Based on the bulbous labellum, the Y-shaped ascoids, the symmetric aedeagus and the number of tenacula (two on each surstylus) the specimen is assigned to the genus Philosepedon. According to Chagas et al. (2009) only eight neotropical species of the genus Philosepedon with two tenacula exist. Philosepedon apozonallii n. sp. differs from the already known neotropical species in the following characters: 1) from Philosepedon helicis Dyar, 1929 (see comment on the generic assignment of this species above) and Philosepedon aliciae IbáñezBernal & Cáceres, 2005, in the shape of the anterior branches of the ascoids, which are foliaceous in the two latter species; 2) from Philosepedon mauroae Wagner & Masteller, 1996, in the wing venation, with R1 strong, and the shape of the surstylus strongly curved in P. mauroae; 3) from Philosepedon tritaxis Quate, 1996, in the rows of facets of the median extension, which are three in the latter species, and the shape of the surstylus, which is short and stocky in P. tritaxis; 4) from Philosepedon vieirai Chagas et al., 2009, in the shape of the scape, which is sub-spherical in P. vieirai, the wing venation, with the medial fork incomplete in P. vieirai, and the shape of the surstylus; 5) from Philosepedon oblongolum Quate, 1996, and Philosepedon uncinatum Quate, 1996, in the oval apical enlargement of R2 and M3 and the wing venation with an incomplete M2 in the latter two. Matuna n. gen. (Plate 2, Figs B, C; Text-fig. 6 A–D) Type species: Matuna bechlyi, original designation. Holotype of the species is a male from the collection of the UCMP with inventory number B-6899-41. Locality of the type species: Early Middle Miocene amber from Simojovel de Allende, Chiapas, Mexico. Etymology: The generic epithet refers to Matun, the Mayan word for amber.

Diagnosis: Philosepedon- or Threticus- like genus which can be morphologically distinguished from all other taxa by the conspicuous shape of the antennal segments. Antenna with 16 antennomeres, scape cylindrical and tube shaped, pedicel long ovoid and not completely separated from scape by a suture (Text-fig. 6 C), flagellomeres nodiform, flagellomeres 1–4 asymmetric, terminal three flagellomeres separated, subequal in size and with internodes, ascoids Y-shaped

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(Text-fig. 6 D); gonostyli slender; surstyli short and stocky, with three long tenacula (Text-fig. 6 B). Remarks: By having the eyes reniform with an eye bridge, the flagellomeres nodiform and R with five branches Matuna n. gen. clearly belongs to the subfamily Psychodinae. The general appearance and morphological characters of the head, like eye bridge with 4 rows of facets, labellum bulbous, 14 flagellomeres and those beyond 11 reduced in size resemble representatives of the genus Philosepedon (sensu Quate 1958, 1996) while the number of tenacula (three on each surstylus) would indicate a relationship with the genus Threticus Eaton, 1904 (Vaillant 1973, Ibáñez-Bernal 2004a). However, the unique and unusual antennal structure, with scape and pedicel not distinctly separated by a suture, does not fit the gener-

ic diagnosis of Philosepedon or Threticus nor of other genera of Psychodinae. From findings of a further poorly preserved fossil in Simojovel amber, Mx 235, with a similar antennal structure, a deformation of the antenna due to fossilisation processes can also be excluded. Matuna bechlyi n. sp. (Plate 2, Figs B, C; Text-fig. 6 A–D) Type material: Holotype is a male from the collection of the UCMP with inventory number B-6899-41. Type locality: Miocene amber from Simojovel de Allende, Chiapas, Mexico. Etymology: The specific epithet is dedicated to Dr. Günter Bechly (State Museum of Natural History in Stuttgart).

Diagnosis: same as for genus.

Text-fig. 6. Matuna bechlyi n. gen. et sp. (A) wing. (B) genitalia, lateral view. (C) antenna: scape, pedicel and flagellomeres 1–2. (D) antenna: flagellomeres 11–14.

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Description male: Head: without vertical bristles; eye bridge with four median facet rows, distance between eyes equals one to two facet diameters. Antenna with 16 antennomeres, 0.9 mm in length; scape cylindrical and slender; pedicel long ovoid, not completely separated from scape by a suture (Text-fig. 6 C). Flagellomeres 1–4 asymmetrical with tubeshaped neck inserted at adverse side of oval node, following nodes with very long and slim necks, flagellomeres 1–10 with nodes and internodes of constant size and length, internodes one and a half times longer than nodes. Flagellomeres 11–13 with short internodes, terminal three flagellomeres (12–14) reduced in size, less than half the length of previous flagellomere, flagellomeres 11 and 12 globular, flagellomere 13 rather oval (Text-fig. 6 D). Flagellomeres 1–11 with very fine Y-shaped ascoids. Maxillary palpus with four segments, length of palpus: 0.3 mm. Relative length of palpal segments: 10 – 16 – 20 – 23. Labellum apparently bulbous. Wing (Text-fig. 6 A): Right wing basally folded, left wing clearly visible, without vestiture, about 1 mm in length. Forks complete; medial fork before centre of wing and well basad of radial fork; radial fork beyond centre of wing. R5 ending in acute wing tip. Sc ending beyond base of R1. Crossvein r–m present.

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Genitalia (Text-fig. 6 B): Gonocoxites cylindrical with basal protuberance, with tuft of long hairs basally; gonostylus slender, with acute apex, slightly bent apically. Surstyli stocky with basal protuberance; about as long as tergum 9, with rounded apex bearing three long clavate tenacula, two terminal and one subterminal in position. Aedeagus not very well visible, distiphallus apparently arrowhead-shaped with bifid apex. Matuna sp. (Text-fig. 7 A, B) Material: Specimen is a male from the collection of the SMNS with inventory number Mx 235. Locality: Miocene amber from Simojovel de Allende, Chiapas, Mexico.

Head: Antenna with 16 antennomeres, scape cylindrical and slender; pedicel long ovoid, not completely separated from scape by a suture (Text-fig. 7 B). Flagellomeres 1–6 asymmetrical with tube-shaped neck inserted at adverse side of ovoid node, following nodes with very long and slim necks. Flagellomeres 11–13 with short internodes, terminal three flagellomeres (12–14) reduced in size, less than half the length of previous flagellomere, flagellomeres 11 and 12 globular, flagellomere 13 rather oval. Flagellomeres with

Text-fig. 7. Matuna sp. (A) genitalia, lateral view. (B) antenna: scape, pedicel and flagellomeres 1–2.

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very fine Y-shaped ascoids. Maxillary palpus with four segments. Labellum not visible. Wing: Forks complete; medial fork before centre of wing and well basad of radial fork; radial fork beyond centre of wing. R5 ending in acute wing tip. Genitalia (Text-fig. 7 A): Gonocoxites cylindrical, with a median swelling; gonostyli slender, with acute apex, bent apically, crossing at midline. Surstyli stocky with basal protuberance and rounded apex, bearing three long clavate tenacula. Remarks: Based on the conspicuous shape of the first two antennal segments, with a cylindrical and slender scape and a long ovoid pedicel that is not completely separated from the scape by a suture, the specimen belongs to the new genus Matuna. The specimen shares with Matuna bechlyi n. sp. the general shape of the antenna. However, the venation of the wing and the distiphallus are not accurately visible in Mx 235. Furthermore the gonocoxites differ in shape from M. bechlyi n. sp. by missing the basal protuberance but having a median swelling. Therefore the specimen cannot be assigned to M. bechlyi n. sp.

4. Discussion Beside systematic descriptions of six new species and one new genus the composition of the psychodid fauna in Mexican amber has been analysed with regard to the ecological significance of the fossils for the reconstruction of the former environment. The most common inclusions in the amber material investigated here belong to the Psychodinae (see Table 1). This is the largest subfamily within the Psychodidae and currently consists of 2000 species in about 100 genera (Wagner et al. 2008). Psychodinae are abundant in the Neotropical region including Mexico today (Quate 1955, 1996, Quate & Brown 2004, Wagner & Ibáñez-Bernal 2009) and they are well represented in Dominican, Baltic, and Mexican ambers (Evenhuis 1994). Psychodinae inhabit a variety of aquatic to terrestrial habitats and ecological requirements may differ greatly on generic level. The genera Psychoda and Philosepedon belong to subaquatic and even terrestrial Psychodinae. Psychoda is a dominant genus and shows a high numerical abundance with a great variety of species today (Duckhouse 1966). Extant species are distributed globally and they are well represented in North America (Quate 1955) as well as in the Neo-

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tropical region where Psychoda is represented by 55 recent species (Cordeiro & Bravo 2008). Larvae of recent Psychoda species are saprobiotic and can be found in human biotopes like drains, toilets and cow and horse dung (Vaillant 1971). Based on the lifestyle of recent representatives it can be concluded that the fossil species inhabited decomposing organic matter on marshy ground near water bodies. The genus Philosepedon has been recorded with 91 species worldwide, and 28 of these are reported from the Neotropical region (Bravo et al. 2006). Because the developing larvae exclusively feed on dead molluscs and snails ( Jung 1956, Wagner & Masteller 1996) it can be expected that the fossil species has had the same feeding preference and was associated with gastropods. Representatives of the genus Threticus are mainly distributed in the Holarctic, Ethiopian and Australasian regions today. The larvae are aquatic to subaquatic. The rheophobe larvae of Threticus aenigmaticus can be found in moist and decaying vegetations in forests (Duckhouse 1966), but most species probably prefer cool fresh water and can be found on slopes with trickling water or even in edges of streams in wet forests like the rheophile larvae of Threticus irroratus Satchell, 1953 (Duckhouse 1966). Hence the fossils indicate the presence of running waters or standing water bodies. With 54 described species the genus Alepia is well represented in the Neotropical region today (Omad & Rossi 2012). The biology of Alepia is poorly known but different authors report the abundance of living species in bromeliads (Duckhouse 1974, Quate & Brown 2004, Wagner & Hribar 2004, Frank & Lounibos 2009) that impound water in their inner leaf axils which provides a biotope for a variety of subaquatic arthropods and other animals. But since Quate & Brown (2004) reported that Alepia longinoi has been found in branches of Cecropia obtusifolia, inhabited by a colony of the ant Azteca, the ecological requirements of Alepia seem to differ greatly on species level as the development of the larvae apparently depends on subaquatic to even terrestrial habitats. The second most common fossils in Mexican amber belong to the Trichomyiinae, which account for almost half of all specimens in the investigated material (see Table 1). The subfamily includes only one extant genus, Trichomyia, with 146 species worldwide and 74 of these distributed in the Neotropical region (Bravo & Araújo 2013). In Mexico only three re-


cent species have been recorded (Ibáñez-Bernal 2004b). Overall 21 fossil taxa of Trichomyiinae are known today (Evenhuis 1994, EDNA Fossil Insect Database, accessed on August 2013: http://edna. palass-hosting.org) and prior to the present study six species of Trichomyia have been described from Mexican amber (Quate 1961, 1963). Adults of recent species can be found resting under bark of trees and show a high abundance in wet rotting woody material derived from tree holes and hollows between buttress roots (Duckhouse 1978). Larvae of Trichomyia are xylophagous or feed on bacteria and fungi (Wagner & Hribar 2004). Since the production of resin is a defence response against a great diversity of phytophagous insects and pathogenic fungi (Langenheim 2003) dying trees are more likely to trap insects like Trichomyia. Therefore, the high amount of Trichomyia in Mexican amber and other amber deposits (Evenhuis 1994, Grund 2006, Wichard et al. 2009) in contrast to other genera of Psychodidae is interpreted as a taphonomic phenomenon. Moreover the dependency on dead wood, especially for breeding, would explain the large proportion of females in the investigated amber material which are represented by 17 specimens in contrast to 3 males. With only two described species, Pintomyia bolontikui Ibáñez-Bernal et al., 2013, and Micropygomyia paterna Quate, 1963 (originally described as Phlebotomus paternus by Quate 1963 but belongs to the genus Micropygomyia according to Andrade Filho et al. 2009a), Phlebotominae are rare in Mexican amber from Simojovel. In contrast, 15 fossil sand fly species have been described from the Miocene Dominican amber (Andrade Filho et al. 2009a, b) with 12 of these belonging to the genus Pintomyia (sensu Galati 2003), and recently three fossil phlebotomines have been found in the comparatively poorly investigated Totolapa amber from Mexico. Females of Phlebotominae are blood feeders and their presence in amber indicates vertebrate hosts like small mammals, reptilians and birds. The larvae develop in humid but not saturated soils, rich in organic matter and protected from desiccation, such as animal burrows, on dead leaves on the forest floor, at the base of trees, in debris and other similar moist microhabitats (pers. com. Ibáñez-Bernal 2013). Taking into consideration the abundance of Phlebotominae in other amber deposits and the ecological requirements of the larvae, their rarity in amber from Simojovel

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could have environmental reasons, such as temporarily saturated soils. However, further studies on all amber deposits from Mexico are needed to investigate this issue. In general the fossil Psychodidae from Mexican amber show various distribution patterns today. Besides cosmopolitan genera like Psychoda, the amber contains characteristic Neotropical elements which are common in South America and Mexico today, such as Alepia and Philosepedon, and primarily Northern Hemisphere elements like the genus Threticus. Because of its location Mexico probably was and still is a faunal interface with elements of both the North American and the South American fauna. The majority of the investigated Psychodid inclusions belong to terrestrial or subaquatic genera that develop in decaying organic matter on the ground or rotting wood today, true aquatic taxa are scarce in Mexican amber. The fossils depict the former environment as a tropical forest with damp to marshy ground, containing large amounts of decomposing organic matter, rotting woody material and water sources in the form of standing water bodies or running waters. New findings of fossil Psychodidae in Totolapa amber from Mexico offer possibilities for future comparative ecological studies. Furthermore, a current project about taphonomy in Mexican amber revealed more than 1000 specimens of recent Psychodidae. A comparison of the fossil faunas from all deposits of Mexican amber with the extant Psychodid fauna from the south of Mexico will contribute to the understanding of former ecosystems and of the development of diversity in a region which is today a megadiversity area. 5. Acknowledgements This research was possible with a postdoctoral fellowship to M.M.S.K, No. SO894/3-1 from the German Research Foundation (DFG) and with an Annette Kade Fellowship to F.S. from the Richard Gilder Graduate School, Museum of Natural History, New York. The authors would like to thank Mark Goodwin from the University of California, Museum of Palaeontology, Berkeley and Dr. Günter Bechly from the Stuttgart State Museum of Natural History Schloss Rosenstein for the loan of the specimens. Special thanks are due to Prof. Dr. Jes Rust (Steinmann Institute, University of Bonn) for his continuous support, Prof. Dr. Rüdiger Wagner (University of Kassel) and Dr. David Grimaldi (American Museum of Natural History, New York) for valuable comments and helpful discussion, Dr. Sergio IbáñezBernal (Instituto de Ecología, Veracruz) for reading and correcting the manuscript, Prof. Dr. Alan Lord (Senckenberg Research Institute and Natural History Museum Frankfurt) for grammar

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correction and Paul Nascimbene (American Museum of Natural History, New York) for technical support.

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Explanation of Plates Plate 1 Fig. A: UCMP, B-7045-38; Psychoda tzotzili n. sp.; scale: 0.5 mm. Fig. B: UCMP, B-8425-92; Psychoda rusti n. sp.; scale: 0.5 mm. Fig. C: UCMP, B-8419-27; Philosepedon apozonallii n. sp.; scale: 1 mm. Fig. D: UCMP, B-7045-38; Psychoda tzotzili n. sp., genitalia, lateral view; scale: 0.1 mm. Fig. E: UCMP, B-8419-27; Philosepedon apozonallii n. sp., genitalia, lateral view; scale: 0.2 mm.

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Plate 2 Fig. A: UCMP, B-7047-32; Trichomyia richardi n. sp.; scale: 1 mm. Fig. B: UCMP, B-6899-41; Matuna bechlyi n. gen. et sp.; scale: 1 mm. Fig. C: UCMP, B-6899-41; Matuna bechlyi n. gen. et sp.; head and basal part of antenna; scale: 0.2 mm. Fig. D: UCMP, B-8416-6; Trichomyia axeli n. sp; scale: 1 mm. Fig. E: UCMP, B-7047-32; Trichomyia richardi n. sp., genitalia, lateral view; scale: 0.2 mm. Fig. F: UCMP, B-8416-6; Trichomyia axeli n. sp., genitalia, lateral view; scale: 0.2 mm.

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