Tecmessa, Burmeister, 1878

St Laurent, Ryan A., Goldstein, Paul Z., Miller, James S., Markee, Amanda, Staude, Hermann S., Kawahara, Akito Y., Miller, Scott E. & Robbins, Robert K., 2023, Phylogenetic systematics, diversification, and biogeography of Cerurinae (Lepidoptera: Notodontidae) and a description of a new genus, Insect Systematics and Diversity 7 (2), pp. 1-25 : 7-10

Tecmessa and Americerura

Tecmessa contains the type species Thosea annulipes Berg, 1878 from Argentina (later moved to Tecmessa by Burmeister (1878)) and Tecmessa elegans from southeastern Brazil ( Berg 1878, Burmeister 1878, Schaus 1901). Tecmessa annulipes and T. elegans are both similar in patterning to cerurines and have simple genitalia, superficially similar to those of some cerurine genera (e.g., Cerura ), see Figs. 8 and 9 . Schintlmeister (2013), in his comprehensive checklist of global Notodontidae and Oenosandridae , transferred all American representatives placed in Cerura since the time of Draudt (1932) to Tecmessa based on these superficial similarities. This classification was followed by Becker (2014) and Miller et al. (2018). However, the transfer of species from Old World endemic Cerura to Tecmessa by Schintlmeister (2013) was erroneous, although these species do not belong in Cerura either. Morphologically, true Tecmessa display the following characters not shared with Cerurinae : narrowly pectinate (not plumose) antennae and somewhat developed ‘beard tufts’ below the haustellum, the latter of which is a derived condition of Heterocampinae ( Miller et al. 2018) . We also note the strong similarity of Tecmessa genitalia (see revised genus diagnosis below) to some Heterocampinae genera, such as Coelodasys Packard ( Miller et al. 2021) . The caterpillar morphology ( Fig. 10 ) and behavior, which are known for both T. annulipes and T. elegans , are also highly distinct from those of Cerurinae . The caterpillars of these two species lack the main larval apomorphies of Cerurinae , namely stemapods and enlarged thoracic segments ( Berg 1878, Oleiro et al. 2011, St Laurent pers. obs., Wheeler pers. comm.); and both feed on Anacardiaceae , not Salicaceae as is typical of Cerurinae ( Biezanko et al. 1974, Oleiro et al. 2011). We also note that these two species have gregarious caterpillars, whereas those of Cerurinae are nearly always solitary in all stages, except for the aposematic caterpillars of Cerurina ( Mulvaney 2021, St Laurent pers. obs.).

We transfer Tecmessa from Cerurinae to Heterocampinae based on our phylogenetic results ( Figs. S1–4 ) and the characters considered apomorphic for Heterocampinae and Cerurinae by Miller et al. (2018). Two additional taxa, following our examination, are also included in Tecmessa on the basis of genitalia and external morphology: the monotypic genus Corania Schaus, 1939 , syn. nov., assigned to Cerurinae in Becker (2014), is hereby synonymized with Tecmessa : Tecmessa pedrana comb. nov. (TL: Argentina). Tecmessa pica (TL: Chile), which was already transferred to Tecmessa by Becker (2014), is highly similar morphologically to T. pedrana and the genitalia of the two species are nearly identical (compare Fig. 9C and D ). Although DNA extraction of these two species failed to provide enough DNA for AHE sequencing, the genitalia provide sufficient evidence for the placement of T. pedrana in Tecmessa and maintenance of Becker’s (2014) assignment of T. pica to Tecmessa . Furthermore, among the series of T. pica , we found specimens intermediate in coloration and patterning between T. pica and T. pedrana , suggesting a close relationship between these two species. So far as is known, true Tecmessa under this concept of the genus are endemic to South America, inhabiting Brazilian Atlantic Forest, grasslands in Brazil and Uruguay, arid regions of Argentina, and Chilean mountain ranges. Habitus and male genitalia of all known true Tecmessa are shown in Figs. 8 and 9 , and caterpillars of two species in Fig. 10 . The two generic synonyms listed under Tecmessa in Becker (2014), Eucerura Schaus, 1901 and Eunaduna Dognin, 1901 , remain synonyms of Tecmessa because the type species of both were examined and determined to be Tecmessa species. Namely, the type of Eucerura , Drymonia pica Butler, 1882 , is Tecmessa pica and the type of Eunaduna , Eunaduna cerurata Dognin, 1901 , is a synonym of Tecmessa annulipes .

Three other species transferred to Tecmessa by Schintlmeister (2013) belong neither in Cerurinae nor in Tecmessa , but to another as yet unnamed heterocampine genus. These species, T. olindata ( Schaus, 1939) , T. gonema ( Schaus, 1905) , and T. laqueata ( Schaus, 1911) are each very similar to one another but together are divergent in morphology ( Figs. 8E, F , 9E ) and phylogenetically ( Figs. S1–4 ) from true Tecmessa . Only T. gonema is included in our analyses, but external and genitalia morphology of these three species are nearly identical. They differ greatly from Cerurinae and from Tecmessa s.s. in that they bear cteniophores, a widespread trait in Heterocampinae which is absent in Cerurinae ( Miller et al. 2018) and Tecmessa s.s. Maintaining our focus on Cerurinae , we do not formally transfer these taxa from Tecmessa at this time, but they will require a new genus to maintain monophyly of Tecmessa and this is the topic of a future work (St Laurent et al. in review).

Below we describe a new genus to accommodate the cerurine contingent of species formally assigned to Tecmessa s.l. and transfer them accordingly. Adult habitus and male genitalia are shown for all of the transferred species in Figs. 11–13 . Description of Americerura gen. nov. ensures that all genera in Cerurinae , and the subfamily itself, are monophyletic. Miller et al. (2018) provide a detailed description of ‘ Tecmessa ’ consisting of the North American species of Americerura , thus we provide a new description here taking into consideration all Neotropical species and excluding the species now established as heterocampines.

Figures

Fig. 8. Tecmessa adult males.A, T. annulipes, type species (NHMUK); B, T. elegans (NHMUK); C, T. pica (USNM); D, T. pedrana comb. nov., primary type (photo V.Verdecia, CMNH);E, 'Tecmessa′ gonema (USNM); F, 'Tecmessa′ laqueata (USNM). Scale bar = 1 cm.

Fig. 9. Male genitalia of Tecmessa.The excised phallus is shown below the ventral aspect of the genitalia and the eighth sternite is shown to the right of each set of genitalia.A,T.annulipes type species, gen. prep. USNMENT01771001 (USNM);B, T. elegans, gen. prep.USNMENT01771030 (USNM); C,T.pica, gen.prep. USNMENT01771031 (USNM); D, T. cf. pedrana comb. nov., gen. prep. USNMEN01771034 (USNM); E, 'Tecmessa′ gonema, gen. prep. 5-12-22:3 (USNM). Scale bar = 1 mm.

Fig. 10. Tecmessa caterpillars.A, T. annulipes, Uruguay (photo J. S. López); B, T. elegans, Argentina (photo G.Wheeler).

Fig. 1. Examples of Cerurinae larvae from nine of the 14 genera: A, Pseudorethona albicans (photo H. Staude); B, Notocerura spiritalis (photo S. Woodhall); C, Hampsonita esmeralda (photo H. Staude); D, Cerurina marshalli (photo L. Mulvaney); E, Furcula furcula (photo W. Bacher); F, Neocerura liturata (photo R. Tawade); G, Americerura scitiscripta (photo R. St Laurent); H, Cerura vinula (photo L. Jonaitis); I, Kamalia australis (photo C. &T.Deane).

Fig.2. Best IQ-TREE result of 100 independent analyses.Absence of bootstrap values means UFBoot ≥ 95 and SH-aLRT ≥ 80.See Fig.S1 for outgroup relationships and full support values. Caterpillar in bottom left is Americerura scitiscripta showing a defense display (photo J. Miller). Adult cerurine moths shown to the right:A, Pararethona argentescens (photo D.Fischer);B, Pseudorethona albicans (photo M. FitzPatrick); C,Cerurella natalensis (photo Q.Grobler); D, Oreocerura dissodectes (photo R. St Laurent, NHMUK); E, Notocerura spiritalis (photo W. Roland); F, Hampsonita cf. esmeralda (photo N. Voaden); G, Cerurina marshalli (photo L. Mulvaney); H, Afrocerura sp. (photo M. FitzPatrick); I, Neoharpyia verbasci (photo F. Romão); J, Furcula furcula (photo G. Kunz); K, Neocerura liturata (photo S. Lamberts); L, Americerura scitiscripta (photo A. Sourakov); M, Kamalia sp. (photo G. Kunz); N, Cerura vinula (photo A. Hardacre).

Fig. 3. Biogeographic reconstruction of Cerurinae in RevBayes using a constrained, time-stratified DEC model. Nodal symbols represent the highest posterior probabilities (PP) for ancestral ranges, the node symbol size reflecting the PP value given in the inset legend. State codes are as follows:A = Africa, R = Nearctic, N = Neotropical,W = West Palearctic, E = East Palearctic, I = Indomalaya, and U = Australasia; with combined states given as a combination of any two codes, which are further denoted on key nodes for clarity.

Fig. 4. Relationships among Notodontidae subfamilies inferred from the present study based on the best of 100 IQ-TREE analyses.The full tree can be found in Fig. S1. Bootstrap values for all branches shown are 100/100 (UFBoot/SH-aLRT) except for few noted nodes.

Fig. 11. Males of each species of Americerura gen. nov. A, A. annulifera comb. nov. (NHMUK); B, A. argynnis comb. nov., holotype of A. argentina syn. nov. (USNM); C, A. argynnis comb. nov. (photo E. Orlandin, DZUP); D, A. rivera comb. nov. (photo E. Orlandin, DZUP); E, A. trigonostigma comb. nov., holotype (USNM); F, A. purusa comb. nov., holotype (photo V. Verdecia, CMNH); G, A. splendens comb. nov. (photo E. Orlandin, DZUP); H, A. bratteata comb. nov. (NHMUK); I, A. dandon comb. nov. (USNM); J, A. grandis comb. nov. (NHMUK); K, A. presidio comb. nov., holotype (USNM);L, A. lancea comb. nov. (USNM);M, A. scitiscripta comb. nov. (USNM); N, A. xicona comb. nov., holotype (USNM);O, A. duonumennia comb. nov. (NHMUK); P, A. candida comb. nov. (MGCL);Q, A. rarata comb. nov. (MGCL); R, A. cf. tehuacana comb. nov. (NHMUK). Scale bar = 1 cm.

Fig. 12. Male genitalia of each species of Americerura gen. nov..The excised phallus is shown below the ventral aspect of the genitalia.A, A. annulifera comb. nov., gen. prep. NHMUK010316656 (NHMUK); B, A. argynnis comb. nov., holotype of A. argentina syn. nov., gen. prep. USNMENT00991441 (USNM); C, A. argynnis comb. nov., paralectotype, gen. prep. 6-23-22:1 (USNM); D, A. rivera comb. nov., gen. prep. USNMENT01771032 (USNM); E, A. splendens comb. nov., gen. prep. USNMENT01771033 (USNM); F, A. trigonostigma comb. nov., holotype, gen. prep. USNMENT00991268 (USNM); G, A. purusa comb. nov., gen. prep. USNMENT01771008 (CMNH); H, A. bratteata comb. nov., gen. prep. USNMENT01771002 (AMNH); I, A. presidio comb. nov., gen. prep. LEP88199 (MGCL); J, A. dandon comb. nov., gen. prep.5-12-22:2 (USNM);K, A. grandis comb. nov., gen. prep.NHMUK010316655 (NHMUK);L, A. lancea comb. nov., gen.prep. 5-12-22:4 (USNM); M, A. duonumennia comb. nov., gen. prep. LEP88198 (MGCL); N, A. scitiscripta comb. nov., gen. prep. LEP33761 (MGCL); O, A. candida comb. nov., gen. prep. 5-12-22:1 (USNM); P, A. xicona comb. nov., holotype, USNM gen. slide 220 (USNM); Q, A. rarata comb. nov., gen. prep. 5-12-22:8 (USNM); R, A. cf. tehuacana comb. nov., gen. prep. NHMUK010316657 (NHMUK). Scale bar = 1 mm.

Fig. 13. Male eighth sternite of each species of Americerura, all sternites correspond to specimens figured in Fig. 12, gray areas around some sternites are composed of membrane from the abdomen.A, A. annulifera comb. nov.; B, A. argynnis comb. nov., holotype of A. argentina syn. nov.; C, A. argynnis comb. nov.; D, A. rivera comb. nov.; E, A. splendens comb. nov.; F, A. trigonostigma comb. nov.; G, A. purusa comb. nov; H, A. bratteata comb. nov.; I, A. dandon comb. nov.; J, A. grandis comb. nov.; K, A. lancea comb. nov.; L, A. scitiscripta comb. nov.; M, A. presidio comb. nov.; N, A. candida comb. nov.; O, A. xicona comb. nov.; P,A. duonumennia comb. nov.; Q, A. rarata comb. nov.; R, A. cf. tehuacana comb. nov.. Scale bar = 1 mm.