HYM. RES. J. Vol. 19(1), 2010, pp. 44-50 Two New Species of the strigatus Species Complex of the Ant Genus Cyphontyrmex (Hymenoptera: Formicidae) from Costa Rica and Panama William P. Mackay and Francisco Serna DepartmentofBiologicalSciences,CentennialMuseum,TheUniversityofTexas,ElPaso,TX79968, USA and Grupo Sistematica de Insectos Agronomia SIA, Museo Entomologico UNAB, Carrera 30 #45-03, Universidad Nacional de Colombia, Bogota, COLOMBIA [email protected]; [email protected] — Abstract. ThestrigatusspeciescomplexisdefinedasthoseworkersandfemalesofCyphomyrmex in which the preocular carina extends back to the vertex, delimiting the lateral margin of a depressedconcavescrobe. Themandibleshave6ormoreteethandthereisasinglemedialpronotal tubercle. The complex was previously reported only in South America, especially southeastern BrasilandnorthernArgentina. TwonewspecieswerefoundinCentralAmerica: C. andersonifrom CostaRica,and C.snellingifromPanama. CyphomyrmexandersoniresemblesC. quehradae,butcanbe separated as the hind femur is longer than the head capsule (shorter in C. quehradae). It can be differentiated from the similar C. hruchi as the mesosomal tubercles are distinct (indistinct in C. bruchi). Cyphomyrmex snellingi has the frontovertexal corners lobate and somewhat projecting posteriad. Itismostsimilarto C.faunulus,butcanbeeasilydistinguished as theanteriormesonotal tubercleisnotmoredevelopedthantheothertubercles (muchlargerthantheothersinC.faunulus). — Resumen. El complejo strigatus del genero Cyphomyrmex se caracteriza porque las hembras y obreraspresentanunacarinapreocularque seextiendeposteriormentehasta elvertexydelimitala margenlateraldeunescroboantenal deprimidoyconcavo. Estashormigasposeenmandibulascon 6 o mas dientes, y un solo tuberculo pronotal mesial. El complejo strigatus se conocia solo de Suramerica, especialmente el sureste de Brasil y norte de Argentina. Dos nuevas especies fueron halladas en America Central: C. andersoni de Costa Rica, y C. snellingi de Panama. Cyphomyrmex andersoni es similar a C. quehradae, pero se diferencia porque el femurposterior es mas largo que la capsula cefalica (mas corto en C. quehradae). A su vez, C. andersoni puede ser diferenciada de C. hruchi porque los tuberculos mesosomales son distinguibles (no distinguibles en C. hruchi). Cyphomyrmex snellingi tiene las esquinas frontovertexales lobosas y algo proyectadas poste- riormente. Estaespecieesm^as similara C.faunulus,delacualpuedeserdistinguidafacilmentepor el tuberculo mesonotal anterior no tan desarrollado. The ant genus Cyphomyrmex belongs to head, which covers most of the head. The the New World fungus growing ants ofthe mesosoma has a series of pairs of blunt tribe Attini, and presently contains 40 tubercles in nearly all species. The first species (Bolton et al. 2007). The genus is opisthogastral tergum* (see glossary in divided into two species complexes, the Serna and Mackay 2010) lacks tubercles. strigatus complex (Kempf 1964) and the Most surfaces are dull and without sculp- rimosus complex (Kempf 1965; Snelling and ture; the hairs are mostly limited to Longino 1992). Cyphomyrmex workers and appressed, often scale-like setae that are females are easily recognized, as the frontal nearly always restricted to the gaster and carinae form a shield on the dorsum of the the head. . Volume 19, Number 1, 2010 45 Most species nest in the soil, in rotten METHODS AND MATERIALS logs and stumps, or in hollow dead twigs. The specimens were examined with a This genus also nests under bark, under Zeiss stereoscope, at 64X, and were mea- moss, and within epiphytic pseudobulbs sured with an ocular micrometer. The (Snelling and Longino 1992). Colonies are abbreviations are as follows: small, probably not exceeding 500 workers (Snelling and Longino 1992). All Cypho- HL Head length, measured in full myrmex species cultivate badisiomycete frontal view, from anterior fungi in the tribe Leucocoprineae. In the margin of medial lobe of cly- C. rimosus group, these fungi grow in a peus to medial posterior mar- yeast form (small masses of unicellular gin of frons HW fungal cells) ratherthaninthemulticellular Head width, measured in full mycelialformtypical for allother attine ant frontal view, maximum width gardens (Schultz et al. 2002; Schultz and excluding eyes (Measured near Brady 2008). posterior point of head) Workers of the strigatus complex can be SL Scape length, excluding con- recognized by the closed antennal scrobe dyle (sometimes with poorly defined margins), EL Eye length, maximumdiameter mandibles with six or more teeth, and with of eye a single medial pronotal tubercle (appar- EW Eye width, maximum width of entlyafusionoftwotubercles). The species eye, perpendicular to EL WL of the strigatus complex were previously Weber's length, a diagonal line considered to be primarily southern South from the top of the anterior American in distribution, although C. edge of the pronotum to the faunulus occurs as far north as Vene- posterior edge of the pos- zuela (Mayhe Nunes and Jaffe 1998). teropropodeal lobes. An unidentified species is found in Co- CI CephaUc index, HW/HL X 100 lombia (Fernandez and Palacio 1995) SI Scape index, SL/HL X 100 and an apparently new species was OI Ocular index, EW/EL X 100 found in Ecuador (Tiputini) by Kari Ryder MCZC Museum of Comparative Zool- Wilkie (http://people.bu.edu/karitr/genus/ ogy, Harvard University cyphom3mnex.html) CWEM Collection of William and In comparison, the workers of the rimo- Emma Mackay, University of sus complex have an open antennal scrobe Texas at El Paso (anteriorly), with the preocular carina curved mesially in front of the eye, and Terms followed by an asterisk are not directed to the posterior corner of defined at the end of this paper and the head, the mandibles have five teeth, explained in the glossary of Serna and Mackay and the pronotum lacks medial tubercles, (2010). or has a pair of tubercles. The species of RESULTS the rimosus complex are widely distribu- ted from the United States to South Cyphomyrmex andersoni new species America. (Figs 1-6) Two new species of the strigatus species — complex were found in Costa Rica and Diagnosis. The worker is a small (total Panama. These new species wiU be in- length about 2.5 mm, n=2) reddish-brown cluded in a key to the species of Cypho- ant. The mandibles have six teeth, the myrmex that can be found at http://www. frontal carinae do not reach the dorsad utep.edu/leb/antgenera.htm. occular suture, the frontovertexal* corners 46 JournalofHymenoptera Research: Festschrift Honoring Roy Snelling are barely extended into auricle-like struc- (length 0.04 mm) and rounded; petiole tures; the pronotum has three angulate enlarged posteriorly, forming dorsal tuber- processes or teeth, including the medial cles as seen in lateral view (length 0.1 mm, process and two lateral processes, together height 0.07 mm) that appears to have two with a pair of posterior swellings, and the lateral tiny bumps; postpetiole with lon- mesonotum has a pair ofconical processes; gitudinal medial depression flanked by the propodeum has a pair ofanterior,blunt two longitudinal ridges; all femora swol- processes and two well-developed angu- len, fore femur with poorly developed lon- late posterior processes; and the posterior gitudinal carina along posteroventral mar- 1/3 of the petiole is raised into a blunt gin, middle femur similar, but carina process that appears bidentate when seen poorly developed, posterior femur with obliquely from above; the postpetiole has well-developed longitudinal carina form- two parallel raised regions on the dorsal ing distinct angle distad about one third surface; and the gaster lacks longitudinal length from body. raised areas. The posterior femur has a Erect hairs absent, except on mandible; distinctive ventral angulate process, fol- hairs on scape and head appressed, hairs lowed distally by a poorly defined carina. on ventral surface of head and anterior The female and male are unknown. margin ofprocoxa subdecumbent, hairs on — Distribution. Known only from the mesosoma, petiole, postpetiole, legs and states of Alajuela and Guanacaste, Costa gaster appressed. Rica. — WorHkeWr measurements (mm).—HL 0.76- AlaTjyupeela,se2ri7esk. NHaonldot8ypkeWwowreksetro,fSCoasntRaamRoicna,, 0.78, 0.64-0.66, SL 0.58-0.60, EL 0.13- 29-vi-6-vii-1999, R. Anderson # 19901, 99-109B, EW WL 0.14, 0.08-0.10 0.90-0.93. Indices: 10°13'30"N; 34°35'30"W (MCZC). — CI 84-85, SI 74-79, OI 65-73. Mandible Additional material examined. Costa Rica, with 6 teeth; anteclypeusbroadly rounded; Guanacaste, Cacao Field Station, 15-ii-1996, R. paraclypeal teeth* spiniform, moderately Anderson # 1—7682 (1 worker CWEM). developed; frontal lobes and frontal ca- Etymology. This new species is named rinae relatively narrow, extending to front- in honor of Robert Anderson, who col- overtexal corner*, forming carina that fuses lected these specimens as well as thou- with posterolateral margin of antennal sands of other interesting specimens. — scrobe, preocular carina continues poste- Discussion. This specieswould keyto C. riorly to form mesial margin of antennal olitorForel in Kempf (1964), found in Brazil scrobe; eyes relatively small, extending and Argentina. Cyphomyrmex undersoni is HW past sides of head; scape relatively short, somewhat larger (HL of C. olitor 0.64, barely reaches frontovertexal comer; pro- 0.56 from Kempf, 1964), has fewer mandib- notum with medial protuberance, 2 lateral, ular teeth (7-8 in C. olitor), the midpronotal conical tubercles and 2 posterolateral lo- tubercle is approximately the same size as bate processes, anteroinfra angle of latero- the lateral tubercles (midpronotal tubercle pronotum developed; 2 angulate conical much smaller than lateral tubercles in C. tubercles on mesonotum (height approxi- olitor), and the propodeal spines are well mately 0.05 mm); anterior margin of developed (poorly developed in C. olitor). dorsopropodeum with 2 broad processes The mandibular teeth are worn and par- (height 0.03 mm), dorsopropodeum* rela- tiallyhiddenbythe clypeus in theholotype mm tively short (0.08 from notopropodeal andbadlyworninthemandibleoftheother groove to highest point of anterior tuber- specimen,but this species appears tobe the cles) posteropropodeum* longer (0.25 mm, only one of the strigatus species complex measured from anterior tubercles to meta- with six mandibular teeth (and as the pleural lobe), propodeal spines small members of the rimosus species complex Volume 19, Number 1, 2010 47 Anterior mesonotal tubercle Midpronotal Propodeal spine tubercle Scape mm 1 I Figs 1-6. Cyphomyrmexandersoniholotypeworker: 1,sideview.2,mesosomaasseenfromabove. 3,headand mandible,frontalviews.4,topofpronotumasseenfromdorsoposteriorview.5,rightposteriorfemurandtibia posterioraspect. 6, antennal scape. allhavefivemandibular,itisapparentlythe Cyphomyrmex snellingi new species onlyspeciesinCyphomyrmexwithsixteeth). (Figs 7-12) — Biology. The two specimens were col- — lected in montane hardwood leaf litter at Diagnosis. The worker is a small (total 1100-1200 melevation and inwetmontane length 2 mm) ferrugineous red specimen. forest litter. The mandibles have seven teeth, the spini- 48 Journalof Hymenoptera Research: Festschrift Honoring RoySnelling form paraclypeal teeth on the clypeus are developed; anterior and posterior mesono- markedly well developed; the frontal lobes tal tubercles moderately well developed do not reach the inner borders of the eyes and approximately same size, anterior (frontal view of head). The frontovertexal tubercle with slighter broader base; dorso- comers form auricle-like structures; the propodeum shorter that posteropropo- scape is short, and does not reach the deum, propodeum without spines or an- posterior margin of the scrobe. The mid gles; subpetiolar tooth sharp and well pronotal process is angulate, the lateral developed, petiole with two distinct, lon- pronotal tubercles are poorly developed. gitudinal lateral lobes, dorsum ofposterior The anterior mesonotal tubercles are con- face extending over anterior part of post- ical and posterior mesonotal tubercles petiole; postpetiole with longitudinal de- approximately the same size. The propo- pressed regionindorsumofnode, outlined deum is rounded posteriorly and without by two elongated elevated areas; dorsal angles or spines. The subpetiolar tooth is surface of gaster flat, bordered laterally by well-developed and sharp, dorsally the slightly elevated longitudinal areas; all petioleextends overthebase ofthe anterior femorawith carinae alongventralposterior part of the postpetiole, which has two border, that on posterior femur more longitudinal elevated regions, the poster- developed and forming lamina. ior margin of the postpetiole is nearly Few erect hairs on mandibles, antecly- straight; the first opisthogastral* tergum is peus and frontal lobes, remainder of hairs without ridges or processes; all femora are simple and appressed, located mostly on swollen ventrally, with carinae, the poster- head and especially gaster. ior femur has a well-developed ventral All surfaces except mandiblular teeth lamina. and anterior edge of clypeus dull. Erect hairs are sparse, present on the — Type series. Holotype worker (MCZC), 1 mandibles, apex of the scape, ventral paratype worker (CWEM), Panama, Cerro surfaces of the legs, ventral and posterior Campana, 950 m, 5-vi-1995, R. Anderson surfaces of the gaster; appressed hairs are #17833. — abundant on the dorsum of the first Etymology. Named in honor of the opisthogastral tergum. memory of Roy Snelling, recalling a pleas- All surfaces dull, exceptthe region along ant visit to the Los Angeles County base of mandibular teeth which is smooth Museum of Natural History in May of and shiny. 2007 where we spent time with Roy, — Distribution. Known only from the type Gordon Snelling, Brian Brown, and Wei- locality in Pan—ama. ping Xie. — Description. Worker measurements Discussion. Cyphomyrmexsnellingiwould (mm): HL 0.71-0.74, HW 0.58, SL 0.48- key to C.faunulus in Kempfs key (1964). It 0.50, EL 0.09-0.10, EW 0.08, WL 0.85-0.86. can be easily distinguished as the anterior Indices: CI 78-81, SI 64-70, OI 82-86. mesonotal tubercle is relatively small, as Mandible with 7 teeth; spiniform paracly- compared to the greatly enlarged anterior peal teeth very well developed (length mesonotal tubercle of C. faunulus (Fig. 13). 0.07 mm), frontal carinae relatively nar- Additionally, the posterodorsal edge of the rowly spaced, not reaching preocular ca- petiole of C.faunulus does not extend over rina which forms mesiad margin of scrobe; theanteriorfaceofthepetioleasitdoesinC. eyes extending past sides of head, with snellingi. Cyphomyrmex faunulus also lacks about 20 ommatidia; scrobe greatly extend- the erect hairs on the frontal lobes. ing posteriorly, forming auricle-like struc- Although it would key to C. faunulus, the tures; scapes not reaching posterior margin twospeciesdonotappeartobemorpholog- of scrobe; tubercles on pronotum poorly ically similar. Volume 19, Number 1, 2010 49 Midpronotal tubercle Auriculate frontovertexal corner Scape Lannina ^ Preocular carina Cyphomyrmex faunulus Paraclypeal tooth Figs 7-12. Cyphomyrmexsnellingiholotypeworker:7,sideview.8,topviewofmesosoma(basedinpartonthe paratype).9,petioleasseenfromabove,postpetioleasseeninanteriorview. 10,postpetioleasseenfromabove. 11,Head. 12,leftfemurasseenfromposteriorview. Fig. 13,Cyphomyrmexfaunulusmesosoma(ReservaDucke, nearManaus,Amazonas,Brasil, LACM). — Biology. The type series was collected strigatus complex uses only mycelium in a leaf litter extraction from a wet cultivars and is probably plesiomorphic montane habitat. and paraphyletic to the rimosus complex (Schultz et al. 2002). DISCUSSION Cyphomyrmex has two centers of species The genus Cyphomyrmex is divided into richness: the rimosus group at about 10° two species complexes, the rimosus com- north (Mayhe-Nunes and Jaffe 1998), plex and the strigatus species complex. The whereas the majority of the species of the 50 Journalof Hymenoptera Research: Festschrift Honoring Roy Snelling strigatus group is restricted to 20°and 30° LITERATURE CITED south (Sanhudo et al. 2007). The strigatus group also lacks species with wide dis- Bolton,B.,G. Alpert,P.Ward,and P.Naskrecki.2007. Bolton's Catalog ofAnts of the World: 1758-2005. tributions (Mayhe-Nunes and Jaffe 1998) as Harvard University Press, available on CD. is found in the rimosus complex. Brandao, C. and A. Mayhe-Nunes. 2001. A new Apparently no new species of the striga- fungus-growing ant genus, Mycetagroicus gen. tus group have been described since With the description of three new species and Kempf's revision (1964), although the comments on the monophyly of the Attini (Hymenoptera: Formicidae). Sociobiology 38: recently described C. muelleri shows simi- 639-665. larities to thestrigatus species complex, and Fernandez, F. and E. Palacio. 1995. Hormigas de along with C. longiscapus, C. costatus and C. Colombia IV: Nuevos registros de generos y wheelerimaybe related (Schultz etal. 2002). especies. Caldasia 17: 587-596. A similar new genus Mycetagroicus, with Kempf,W.W. 1964.ArevisionoftheNeotropicalants three new species has been recently de- ofthegenusCyphomyrmexMayr. PartI. Groupof strigatusMayr(Hym. Formicidae). Studia Entomo- scribed (Brandao and Mayhe-Nunes 2001). logica 7: 1^4. GLOSSARY . 1965. A revision of the Neotropical fungus- growing ants of the genus Cyphomyrmex Mayr. Anteclypeus (=''apron''): the anterior Part II: group of rimosus (Spinola) (Hym. For- portion of the clypeus attached to the micidae). Studia Entomologica 8: 161-200. Mayhe-Nunes, A. and K. Jaffe. 1998. On thebiogeog- labrum. Dorsopropodeum: the dorsal sur- raphy of Attini (Hymenoptera: Formicidae). face of the propodeum. Ecotropicos 11: 45-54. Frontovertexal comer: the posterolateral Sanhudo,C, A. Mayhe-Nunes,and C. Brandao. 2007. angle between frons and vertex. Quern sao as Cyphomyrmex (Myrmicinae: Attini). Notopropodeal groove: (= "metanotal Biologico 69: 433-434. groove") a transverse groove on the noto- Schultz, T. and S. Brady. 2008. Major evolution- ary transitions in ant agriculture. Proceedings propodeal fusion. of the National Academy of Sciences 105: 5435- Notopropodeal fusion: In workers, the 5440. tergal fusion of the thoracic notum and the , S. Solomon, U. Mueller, P. Villesen, J. propodeum. Boomsma, R. Adams, and B. Norden. 2002. Opisthogaster (adj. opisthogastral): Cryptic speciation in the fungus-growing ants Cyphomyrmex longiscapus Weber and Cyphomyr- (= "gaster") Abd IV to pygidium. mex muelleri Schultz and Solomon, new species Paraclypeal teeth: (="parafrontal teeth" (Formicidae, Attini). Insectes Sociaux 49: 331- -Kempf1964, 1965),referringtotheanterior 343. teeth-like processes on the clypeus. Serna, F. and W. Mackay. 2010. A descriptive morphology of the ant genus Procryptocerus ACKNOWLEDGMENTS (Hymenoptera: Formicidae). Journal of Insect Science (in press). We would like to thank Roy Snelling and Weiping Snelling, R. R. and J. T. Longino. 1992. Revisionary XiefortheloanofspecimensofCyphomyrmexfaunulus. notes on the fungus-growing ants of the genus Two anonymous reviewers provided important com- Cyphomyrmex, rimosus group (Hymenoptera: For- ments. The research was supported by a grant from micidae:Attini).Pp.479-494in:Quintero,D.,and the National Geographic Society to Robert Anderson A. Aiello eds. Insects ofPanama and Mesoamerica: and the Ernst Mayr Fund of the Museum of selected studies. Oxford University Press, Oxford. Comparative Zoology. 692 pp.