Taxonomic position of two fossil social bees (Apidae)

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(1)Rev. BioL Trop., 24(1 ) : 3543, 1 976. Taxonomic position of two fossil social bees (Apidae) by Warwick Estevarn Kerr * and Rubens Alves da Cunha * * (Received for publication September 1 1 , 1975). Abstraet: Numerical taxonomie methods were used to localize two fossil bees: Meliponorytes devíctus and Electrapís proava. The results indicate that both bees belong to the tribe Meliponini (Apidae). M. devíctus was eonfrrmed as Tetragona devícta. E. proava is strongly associated to the superior trigonas (Trígona, Scaptotrígona, Oxytrigona) . and according to the different existing taxonomic schools it should be renamed either Trígona (Roussyana) proava or simply Roussyana proava.. In 1962 one of us (WEK) had the opportunity to examine and take sorne rneasurernents of two fossils in the British Museurn of Natural History. About 70 characters of each bee were rneasured, but due to the position of the fossils in the arnber, only 51 were coincident. Nurnerical taxonornic techniques were used to assess the taxonornic position of these two fossils.. MATERIAL AND METHODS The two fossils rneasured were Electrapis proava (Oligocene, Baltic) and M elipon orytes devictus (Miocene, Burrna). The latter has been renamed Trigona ( Tetragona) devicta (Cockerell) according to Karr & Maule ( I 964). Both are in the collection of the British Museurn of Natural History. The other bees are deposited in the collection of the Departrnent of Genetics, Faculdade de Medicina de Ribeirao Preto, Bra,zil. Both fossils and present species studied are in the following listo. * **. Depto. de Genética, Faeuldade de Medicina de · Ribeirao Preto, 1 4 . 000, Ribeirao Preto, S.P., Brasil. Depto. de Estatística, Faculdade de Filosofía, Ciencias e Letras de Rio Claro, 1 3.500, Rio Claro, S.P., Brasil 35.

(2) REVISTA DE BIOLOGIA TROPICAL. 36. fossi1 species:. Apidade 01.. 02.. TrigoNZ (TetrflgOF/Il) devicta (Cockerell) Electrapis proalltl Cockerell. Present species Apidae 03.. 04. OS.. 06. 07. 08.. 09. 1 0.. 11. 1 2. 1 3. 14. 15.. TngoF/ll (TetragoF/ll) ÍTidipennis Smith (6) MelipoNZ quadrifasciata Lepeletier ( 1 ) Meliporuz scuteJlIlris Latreille (2) MelipoF/ll puncticollis Friese (3) ScaptotrigoF/ll postica (LatreiUe), a male, '(4) ScaptotrigoF/ll tubiba (Smith) ( 1 ) OxytrigoF/ll tataÍTa (Smith) (1) TrigoF/ll (Trigona) hyaliF/ll ta (Lepeletier) (4) Meliponula bocandei (Spinola) (S) Apis mellifera Linné (4) Bombus (Fervidobombus) atratus Franklin (6) Eulaema (Apeulaema) nigrita Lepeletier (4) Euglossa (Euglossa) cordata Linné (4). Anthophoridae. 16. 17. 1 8.. 19.. Xylocopa (Shoenherria) muscaria (Fabricius) (4) Centris (Hemisiella) tarsata Smith (4) Epicharis (Epicharitides) cockerelli Friese (4) Melitoma segmentaria (Fabricius) (4) Megachilidae. 20. . 21.. Megachile (Leptorachis) paulistana Schrotkyi Anthidium (A nthidium) manicatum (Linné) (6) Halictidae. 22.. Augochloropsis cupreola Cockerell (4) Andrenidae. 23.. Oxaea flavescens (Klug) (4). The numbers in parentheses refer to the location of the specimens: ( I ) Pocinhos do Rio Verde, M.G. ; (2) Riachuelo, S.E. ; (3) Obidos, P. A. ; (4) Rio Claro, S.P. ; (5) Luanda, Angola (Africa); (6) Serra Negra, S.P.; (6) British Museum of Natural History 2-94; Bingham CoHection 96-3, Tenasserim, Burma.� Except for the bees numbered 01, 02, 03, and 1 1 , the others are from Brazil. Fifty-one characters were measured under a Zeiss Stereoscope and are shown in TabIe l . Eight bees, numbers 16 to 23 beIonging to the distantly reIated families Anthophoridae, Megachilidae, Halictidae and Andrenidae were included in the anaIysis as "taxonomic indicators". If the fossil species cluster with sorne of a family that is not Apidae, it wouId mean that the similarity coefficients reached low values, that is, distantly reIated..

(3) KERR & CUNHA: Taxonomy of fossil social bees. TABLE 1 List of characters used in the analysis offossil bees by the numerical taxonomic method 01. 02. 03. 04. 05. 06. 07. 08. 09. 1 0. 1 1. 12. 1 3. 14. 15. 16. 17. 18. 19. 20. 21. 22. 23. 24. 25. 26. 27. 28. 29. 30. 31. 32. 33. 34. 35. 36. 37. 3 8. 39. 40. 41. 42. 43. 44. 45. 46. 47. 48. 49. 50. 5 1.. (1) (lI) (I1I). Length of head Width of head Length oC clypeus Maximum width oC clypeus Length oC eye Width oC eye Diameter oC median ocellum Distance between median and marginal ocellum Distance from median ocellum to eye margin Diameter of antennal basis Distance from antennal basis to eye margin Distance from antennal basis to posterior head margin Distance between antennal basis Distance from upper eye margin to head margin I nner distance between upper margin of eyes Length oC malar area Length of mandible Width of mandible basis Width o f mandible apex Length oC first segment of flagellum Diameter o f fourth segment o f flagellum Length of forewing Width of forewing Length of pterostigma Width of pterostigma Length oC marginal cell Length of free segment of radius Number of submarginal cells Number of hamulli Length of thorax Width o f thorax Length of scutum Width of scutum Length of s<;utellum Width of scutellum Length oC tibia (IH) Width of tibia (III) Length of the first tarsal segment (III) Width of the first tarsal segment (IH) Length of tibia (H) Width o f tibia (H) Length of the fust tarsal segment (Il) Width of the first tarsal segment (11) Length of tibia (1). Width of tibia (I) Length of the first tarsal segment (1) Width of the first tarsal �egment (I) Types of hairs on tibia (In) Types oC pollen carriers Length of corbicula Maximum width of abdomen. First pair of legs Second pair of legs Third pair of legs. 37.

(4) 38. REVISTA DE BIOLOGIA TROPICAL. Gower's coefficient of sirnilarity was calculated in order to have, a sirnilarity matrix among operational taxonomic units (OTU's) and this matrix used to construct dendrograms illustrating the relationship among OTU's, Four different clustering techniques were used in the construction of the dendrograms: Weighted Average Linkage; Unweighted Average Linkage; Single Linkage; and a Weighted Pair-Group method using arithmetic average s, Cophenetic correlations were also obtained (Sneath & Sokal, 1973). Part of the analysis was performed using programs written by M.J. Sakin, Medical Research Council, University of Leicester, and implemented to the set of programs in Numerical Taxonomy by A.M.F. Cadete of the Centro de Cálculo Científico, of the Gulbenkian Science Foundation, Oeiras, Portugal. RESULTS The results are summarized in Fig. 1 (Weighted Average Linkage), Fig. 2 (Unweighted Average Linkage), Fig. 3 (Single Linkage) and table 2 (Reduced Matrix of Similarity Coefficients). TABLE 2. Matrix of similarity coefficients for 1 0 selected OTU's those in the initial similarity matrix) OTU No 02 03 05 06 07 08 09 10 11. (The similarity values are the some as. 01. 02. 03. 05. 06. 07. 08. 09. 10. .758 .947 .649 .682 .814 .764 .843 .745 .707. .757 .652 .695 .818 . 7 74 .853 .771 . 702. .782 .806 .816 .822 .840 .821 . 7 95. .869 .752 .809 .759 .796 .854. .840 .888 .808 .862 .872. .899 . 879 .837 .807. .863 .882 .863. .862 .818. .837. Fig. 1 shows that the group with the species Apis mellifera ( I 2), Euglossa corda¡p ( I 5), Melitoma segmentaria (I9), Megachile paulistana (20), Centris tar sata (I7), Epicharis cockerelli ( I 8), Bombus atratus (I3), Xylocopa m uscaria (16), Anthidium manicatum (2 1 ). Oxaea flavescens (23) and Eulaema nigrita ( 1 4) should be discarded for two reasons: it did not contain any of two fossil bees; and, on the other hand, it contained 7 of the 8 "indicators". In tlús ca!Oe the group: Trigona devicta (O 1 ), Tdgona iridipennis (03), Electrapis proava (02), Melipona quadrifasciata (04), Melipona scutellaris (OS), Melipona puncticollis (06), Melipo nula bocandei ( I 1), Scaptotrigona postica (07), Scaptotrigona tubiba (08), Augo chloropsis cupreola (22), Oxytrigona tataira (09), and Trigona hyalinata ( I O) is left for futher analysis. From Fig. 2 the following group, not discarded, contains bees: 01, 02, 03, 07, 08, 09, 10 and one "indicator" (22). Finally, in Fig. 3 the non discarded group contains the be es 01, 03, 04, 05, 06, 07, 08, 1 0, 09, 1 2, l l , 15, 02 plus the "indicators" 22 and 19. Taking this information into account ethe species in Table 2, were clustered using the "Weighted Pair-Group Method" (Sneath & Sokal, 1973). Fig. 4 shows the dendrogram obtained by this method; it gives the final position of both fossils, Tetragona devicta and Electrapis proava..

(5) KERR & CUNHA: Taxonomy o� fossil social bees. 39. DISCUSSION According to Nelson ( 1 972) the systematist faces three positions: a) the classification should clos�ly express the group phylogeny; b) the classification is an artifice, useful for the phylogenetic studies and; c) phylogeny and classification should be undertaken independently. Our position is the second, especially in this case where we are trying to determine the positon of two fossil bees. The final result of this study (Fig. 4) confirms Kerr and Maule ( 1 964) as far as the position of the Miocene fossil Tetragona devicta is concemed, that is, that it is very close to the present Tetragona iridipennis which still inhabits the same area. The similarity coefficient between these two species is 0.947. Electrapis proava possesses the greatest similarity coefficient with Oxytrigona tataira (0.840) followed by Scaptotrigona tubiba (0.822) and T. hyalinata (0.82 1 ). The two species of Melipona have among themselves a coefficient of similarity of ()'869 , and the two of Scaptotrigona a coefficient of similarity of 0.899. These va!ues suggest that Electrapis proava is a Meliponini associated to superior South American Trigonas, all of which possess communication by trail marking. Manning ( 1 960) divided the fossil genus Electrapis in three subgenera: Electrapis (Electrapis) Cockerell ( 1 909), Electrapis (Protobombus) Cockerell ( 1 909), and Electrapis (Roussyana) Manning ( 1 960) to which E. proava .belongs. Since Manning attributes Apis-like characters to the first subgenus and Bombus-like body and ApiPlike venation to the second, it is obvious that after the present results E. proava should be included in the Meliponini and ca!led Trigona (Roussyana) proava, according to the classification of Moure (Wille & Michener, 1 973). Probably the Oligocene fossil Apis palmnickenensis Roussy, that is considered by Manning ( 1 960) as belonging to the subgenus Roussyana, is also a Trigona (Roussyana) palmnickenensis. This conjecture is supported by plate 1 3 of Roussy ( 1 93 7) which shows that Apis palmnickenensis has many Trigona-like characteristics.. ACKNOWLEDGMENTS This paper received financia! support from the Rockefeller Foundation and the State of Sao Paulo Research F9undation (F APESP). Acknowledgements are also due to the British Museum of Natural History, to the Gulbenkian Science Foundation, and to Dr. Julian Adams who corrected the English version.. RESUMEN Los resultados de lÍbicación de dos abejas fósiles, usando métodos de taxonomía numérica, indican que ambas, Meliponorytes devictus y Electrapis proa va, pertenecen a la tribu Meliponini (Apidae). Se confirmó a M devictus como Tetragona devicta. Por su fuerte asociación con las trigonas superiores ( Trigona, Scaptotrigona, Oxytrigona) y de acuerdo con las diferentes escuelas taXonómicas existentes, E proava debería llamarse Trigona (Roussyana) proava, o simplemente Roussyana proava..

(6) 40. REVISTA DE BIOLOGIATROPICAL. LITERATURE ClTED Kerr, W. E., & Vilma Maule 1 964. Geographic distribution of stingless bees and üts implications. J. New York. Entorno/. Soc., 72: 2- 17. Manning, F. 1 96 0. A new fossil bee from Ba/tic amber. Verh. XI Kongr. Ent. Wien 1 960, Vienna: 306-308 pI. 5. Nelson, G. J. 1 972. Phylogenetic relationship and classification. Syst. Zoo/., 2 1 : 227-2 3 1 . Roussy, L. 1 9 7 3 . Contributions a l'étude de l'abeille tertiaire, de ses parasites et de ses enernis. Gaz. A pico/e, 3 8 : 49-72. Sneath, P. H. A., & R. R. Sokal 1973. Numerical taxonomy. W. H. Freernan, San Francisco. Wille, A., & C. D. Michener 1973. The nest architecture of stingless bees with special reference to those of Costa Rica (Hyrnenoptera: Apidae) . Rev. Biol. Trop., 2 1 (Supl. 1 ) : 1-278.. Fig. 1.. Diagrarn of relationship for the i3 OTU's obtained by the Weighted Average Linkage rnethod. Cophenetic correlation : r = 0.5 2 .. Fig. 2.. Diagrarn of relationships for the 23 OUT's obtained by the ·Unweighted Average Linkage rnethod. Cophenetic correlation: r = 0.73.. 'el.

(7) 41. KERR & CUNHA: Taxonomy of fossil social bees. 0,8. 0,7. 0,6. 0,5. 0,4. 0 ,3. 0,9 01 03. ,-______. 02 04 05. _ _ _ 06 L-. 1 1 07 08 22. ,-____ L-____. 09 10. r------. 12. L-_____. 19. L_ _ _ _ 1 5. _ 20 _ _ _ _ _ '--. '_ _ _ _ _ _ 1 7 '_ _ _ _ _ _ _ _ 1 8. ,-_______ ,-______. 13 16. _ _ _ _ _ _ 21 '-. _ _ _ _ _ _ _ _ _ _ _ 23 L-. '-______________. 14. �gona ( Telragano ) deviclo ( Cockere" ) �gono ( Telrogono ) iridi�ennis Smilh Electra� proova Cockere t l. M . l i � guodrifasciata Lepelefier Melipona � Lat,eille Melipona �uncticollil Fries. Moliponulo bocondei ( Spinolo ) Scoptotrigono postico ( Lotreille) t o mole 9 Scoptotrigono lubibo, ( Smith ) Augochloropsis �preolo Caell,e,el l Q!<ytrigono toloiro ( Smith ) !!!g ono (!!!g ono ) �yolinoto ( Lepeletier ) Apis m e l l ifera Linné �-;o ..o (�gloS$ o ) cordato Unní Melilomo !!9mentorio ( Fobricius) Megochile (Leptarochis) poulistono Schrotkyi. Centris (He;;;;-siello) torsoto Smith Epichoris �itid .. ) cockere " i Friese ;ombus ( F�vidobombus) � Fronklin Xylacopo (Shoenherri o ) muscaria ( Fobricius) ) Anthid ium ( A n thidi u m ) �um ( Linne' . Oxoeo flovesc ens ( Klug ) Euloemo ( �'pe u l o em o ) !!!.grito Lepeletier. 1 0,5. 0,6. 0,7. 0,8. 0,9 01 03. ,-______ '--___. 1----. _ _ _. 02 07 08 22 09 10 04 05. '----- 06 _ _ _ _ 1 1 1-. ,-____ '-____. 12 15. ,------ 1 9. '-_____ '-_______. ,--_______ ,-______ '-______ '-________. 20 17 13 16 21 18. _ _ _ _ _ _ _ _ _ _ 23 L-. 14. Trigono (Tetrogono) deviclo ( Cockere l l ) �oona (Tetragona) i ridipennis Smith Electropis proovo Cocke r e l l scaPtot�ono postico ( Latrei I l e), o mole Scoptotrigono tubi bo, ( Smith ) �gochIO'OPl!! �preola Cockerell Oxytrigono tato ira (Smith ) :¡:;;gon.;-(Trigon o ) �yolin ato ( Lepelelier) MeJipono 9uodrifosciota Lepelefier Melipona scutellaris Latreille Melipona punctico l l i s Fries. Melipan ulo bo condei ( Spinolo) A p i s mel l ifera Linne# Eu;osso ( E u gloss a ) cordato Linné M elitomo .egmentoria ( Fa brici u s ) Mogochi l o (LePtorochi') poulistana Schrotkyi Cen�H.��orsot. Smith 8om bu, ( Fervi dobombu,) atrotus Fro n k l i n X y l o c o pa ( Sh o enherrio) musca ria ( Fobricius ) Anthidi;;;;' (Anlhidi u m ) m� ( U nn') �p i c h o ris ( �.Pi cha ritides) c o c k . r . " i Frie .. Olooa tlo.lSconl ( 1< Iug ) E u l . . .. a ( �p.ulo.mo) �grito. 2. Lopeloti or.

(8) 42. REVISTA D E BIOLOGIA TROPICAL. Fig. 3.. Diagram of relationships for the OTU's obtained by the Single Linkage method. Cophenetic correlation : r = 0.78.. F ig. 4.. Diagram of relationships for 1 0 selected OTU's obtained by the Weighted Pair Group method using aritmethic averages..

(9) 0,6. 0,7. 0 ,8. 0,9 0 1 Trigona (Talrogono ) daviclo ( Cockare l l ) 03. !;gona. (Tatragona ). iridipennis. 04 Mel i pona 9ua drifa sciata. Smilh. Le p e l , I i ... 0 5 M e l i pona s c u l o l l a r i s Lat"i l l . 06 Melipana puncti co l l i s Friase 07. Scap lotrigona. 22. A u go c h l a r o p s i s. 0 8 Scap totrigona. 11-----. poslica ( Lalrei l l el t u b i ba , ( S m i t h ) cupreola. a. malo,. Cockero l l. 1 0 Tri gono ( Tr i ll on o ) !!.yo l i n a ta ( Le p e l o t i o , ) 09 �Jtr i g o n a tola i r a l S m i t h ) 1 2 �p� � Linné �ponulo. 11. bocandoi ( Spinola). 1 5 E u g l o s s a ( E u g l o s . a ) cord a t a L i n n é 1 9 M e lito m a s a g ;;;;,;ta ria ( Fa bri c i u s ). 02 E l e ctra p i . 20 Megachile 17 16. L--_ _ _ _ _ _. proava. Cen�Hemi.. �y l o co �. C o c k e re l l. ( Le p t o ra c h i . ). p a u l i.tana. i�a rsata. ( S h o e n h erria ). S m ith. S c h rotkyi. m u s c a ria ( Fo br i c i u s ). 2 1 A n t h i d i u m ( A n f h i d i u m ) m a n i c a l u m ( Li nn.) 1 8 E pic h a r is ( E p i c h a r i l i des ) c o ck a r e l l i Friasa 13 23 14. Bombus. ( Fe r vido bo m b u s ). � tlavascans E u l aamo. ( KIU9). (�pe u l o em a ). o lralus. �g�. Fron k l i n. L a p e l etier. 3. 0, 700. 0,800. 0,900. 1 , 000 03 Trigono 0 1 Trigona. '-_ _ _ _. 02 07 08 09. ( Tetrogono ). ( Telrogono ). E l actropis. proovo. Sco ptotrigono. Scoptot rigona. iridipennis. Smilh. devicto ( Co ckerel l ). Cocke re l l. postica ( L otra i l l e), tUbi b o , ( S m i th ). Oxytri gono. tatoiro. Melipo n u l o. b o c o n d e i ( S pino l o ). ( Smith ). o mole. 1 0 Tri g o n ;(Trigona ) �yol inata ( Lepeletier ) 05 M e l i p o n o s c u le l l a r is L o t re i l l e ,----- 06 !!!!!ip ono pu nclico l l ;S Friese 11. 4.

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