Return to the Subspecies Concept in the Genus Zeta (Hy- Menoptera: Vespidae; Eumeninae )

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Return to the Subspecies Concept in the Genus Zeta (Hy- Menoptera: Vespidae; Eumeninae ) Bol. Mus. Nac. Hist. Nat. Parag. Vol. 14 (1-2), Setiembre 2002, pp. 19 - 24 RETURN TO THE SUBSPECIES CONCEPT IN THE GENUS ZETA (HY- MENOPTERA: VESPIDAE; EUMENINAE ) JAMES M. CARPENTER Division of Invertebrate Zoology, American Museum of Natural History, Central Park West at 79th Street, New York, NY 10024, U. S. A. e-mail: [email protected] Abstract.- The subspecies recognized in the eumenine wasp genus Zeta are confirmed as synonyms. Resumen.- Las subespecies reconocidas en el género de avispa eumenina Zeta son inveteradas como sinónimos. In 1987, I published a study of the subspe- Ma che dire allora delle 16 sottospecie cies of two species of the neotropical potter dell’‘Eumenes flavopictus (Blanch.)’ wasp genus Zeta. At that time, the four recog- riconosciute e descritte del Van der Vecht nized species in Zeta, were divided into 15 sub- (1959, Zool. Verh. Leiden, 41:1-71); o delle species, with most of the subspecies proposed 15 sottospecie del Polistes rothney Cam., pure studiate dal Van der Vecht (1978, by Giordani Soika (1975). I documented that Bijdr. Dierk., 38/97-109); o delle 17 the color characteristics by which most of the sottospecie di Polistes stigma (F.) del subspecies were distinguished: (1) showed dis- Petersen (1987, Ent. Scand., 10:227-259); crepancies between actual specimens and o delle 9 ‘color forms’ dell’Eumenes Giordani Soika’s (1975) key, (2) were transi- lepeleterii Sauss.» del Bequaert; od anche tional. Distributional overlap was also noted. le 10 «color varieties» dell’Eumenes I concluded that these subspecies were not re- maxillosus (DG), pure del Bequaert (1918, ally distinct, and sank most of them. I also Bull. Am. Mus. nat. Hist., 39: 276-278, e opined that “Recognition of subspecies is in 60: 279-281). È però opportuno ricordare my view a poor way of dealing with color varia- che molte delle «color varieties» del Bequaert sono oggi considerate buone tion in vespids anyway” (Carpenter, 1987: specie. E non è impossibile che qualche 257). sottospecie dell’argillaceum si riveli, ad un Giordani Soika (1990: 166-170) took issue più attento studio, anch’essa valida specie. with this conclusion, in an appendix to a pub- Posso già dire che lo studio dell’apparato lication dealing with some other neotropical copulatore di alcune ha già messo in eumenine genera. Below I quote the full ap- evidenza significative differenze, pendix: specialmente nei lobi basali dell’edeago e negli apodemi, come risulta dall’esempio “In una recente pubblicazione (1987, delle figg. 8991: le differenze tra le PSICHE [sic], 94:253-259) il Carpenter sottospecie incarum ed apurimacense enuncia ‘A review of the subspecies con- dell’argillaceum non sono minori di quelle cept in the eumenine genus Zeta’, basata dalla specie mendozanum (Schr.). sullo studio tassonomico di alcune specie Il Carpenter ritiene fonte di difficoltà di questo piccolo genere neotropicale. Il un ‘couplet’ della mia Tabella per le Carpenter si mostra altamente meravigliato sottospecie dell’argillaceum, che definisce dal fatto che alla specie argillaceum (L.) ‘vague’, e così traduce; vengono attribuite 10 sottospecie: ‘Giordani ‘Ferrugineous, or brown-ferrugineous Soika recognizes only four species in Zeta, and black, without yellow markings. In but these are divided into no fewer that 15 some exemples (transitional to hubrichi) subspecies, 10 of which are in argillaceus!’. part of the posterior margin of the pronotum 20 BOL. MUS. NAC. HIST. NAT. PARAG. VOL. 14 (1-2) and apex of tergum 1 may be yellow’. avvenne, e gli sperimentatori, del tutto ‘Yellow markings fairly extensive both stupiti, conclusero per la non trasmissibilità on thorax and abdomen’. ereditaria dei caratteri acquisiti, e ne fecero A dire il vero, questo ‘couplet’ mi una pubblicazione. sembra assai più chiaro di molti di cui Oggi noi potremmo ritornare al concetto abbonda la letteratura: mi limito a citare - linneano: ‘Species tot sunt quot diversas come esempio - parte della Tabella per la formas ab initio produxit Infinitum Ens; determinazione delle specie di quae deinde formae secondum generationis Stenodynerus (=Nannodynerus Bl.) nella inditas leges produxere plures, at sibi sem- fondamentale Monografia del Bliithgen, per simi les, ut species nunc nobis non sint pubblicata dall’Accademia delle Scienze di plures, quam quae fuere ab initio’. (C. Berlino (Die Faltenwespen Mitteleuropas, Linnaei, Genera Plantarum, Ed. V, 1754, 1961, Abh. Deutsch. Ak. Wiss. Berlin, Kl. p. III). Chem. Geol. Biol., Jh. 1961, 2: 109-110). Ma è oggi difficile accettarlo, e ‘Flügenschuppen zitronen-, hell- oder dobbiamo ammettere la speciazione, e la weisslinchgelb, mit nicht pigmentiertem, razzazione che ne è l’inevitabile fase durchsichtig gelblichem Mittelfleck, und intermedia. E se diamo un nome alle specie linearem solchen Saum, am Innerrand dobbiamo, per chiarezza, precisione, ed geschwàrzt; ausnahmsweise sind sie anche per semplicità, dare un nome anche braunschwarz, mit oder ohne schmale gelbe alle sottospecie. Ovviamente quando le Randbinde, aber nie gerötet’. caratteristiche delle sottospecie sono ‘Flügelschuppen rot oder orange oder nettamente al di fuori dei limiti di hellgelb, mit durchsichtiger roter variabilità della forma nominale o delle iiberlagerung’. altre sottospecie, e sono geneticamente Il nuovo concetto di sottospecie del Car- stabilizzate, sì da ripresentarsi eguali nelle penter è imperniato sul non riconoscimento successive generazioni. della validità delle sottospecie, che A conferma del punto di vista del Car- sarebbero ‘artificial taxa’. Questo concetto penter interviene Mud D’aub (1988, è ripreso dal Carpenter in SPHECOS (1988, SPHECOS, n° 17:12): ‘The use of subspe- n° 17:12), ove approva l’eliminazione delle cies is akin to putting stamps into albums sottospecie proposta dall’Archer ‘especially that have rectangles for each issue’. Non è to the extent that subspecific taxa are elimi- forse tanto pertinente il confronto con nated’. E rafforza questo principio insetti che, certamente, non hanno caselle ricordando le ricerche di McLean (in realtà predisposte; comunque siamo nel più pieno precedute da molte altre di vari AA., concetto linneano «Species tot sunt...’, ove specialmente su Lepidotteri) che hanno 1'«Infinitum Ens» è l’uomo, che stampa gli ‘even shown experimentally that such ‘sub- immutabili francobolli. species’ in Polistes can be produced by Riservandomi di discutere più manipulation of humidity!». ampiamente la questione in un lavoro di In realtà, il fatto che variando alcuni prossima pubblicazione, confermo la fattori ambientali si inducono variazioni di validità delle sottospecie da me segnalate colore in molti insetti, è del tutto ininfluente nella mia nota del 1975 sul genere Zeta nei casi in esame, in quanto tali variazioni Sauss.” sono limitate alla generazione che li ha subiti, e scompaiono completamente già which I translate (after correcting a few ty- alla prima generazione successiva. Ciò mi pographical errors) as: ricorda le ricerche, riportate dai vecchi testi universitari di Zoologia, di alcuni “In a recent publication (1987, Psyche, ‘scienziati’ che, avendo tagliata la coda ad 94:253-259) Carpenter enunciates ‘A re- alcuni topi, si attendevano che la loro prole view of the subspecies concept in the nascesse priva di coda. Questo non eumenine genus Zeta’, based on taxonomic SETIEMBRE 2002 RETURN TO THE SUBSPECIES CONCEPT IN THE GENUS ZETA 21 study of some species of this small literature abounds: I confine to quote - as neotropical genus. Carpenter marvels example - part of the key for the determi- highly at the fact that to the species nation of the species of Stenodynerus argillaceum (L.) 10 subspecies are attrib- (=Nannodynerus BI.) in the fundamental uted: ‘Giordani Soika recognizes only four monograph of Blüthgen, published by the species in Zeta, but these are divided into Academy of Sciences in Berlin (Die no fewer that 15 subspecies, 10 of which Faltenwespen Mitteleuropas, 1961, Abh. are in argillaceus!’. Deutsch. Ak. Wiss. Berlin, Kl. Chem. Geol. But is what one to say then of the 16 Biol., Jh. 1961, 2: 109-110). subspecies of ‘Eumenes flavopictus ‘Tegulae lemon -, brightly - or whitish- (Blanch.)’ recognized and described by van yellow, with not pigmented, lucidly sallow der Vecht (1959, Zool. Verh. Leiden, 41:1- middle-stain, and linear such margin, at the 71); or of the 15 subspecies of Polistes inner-edge blackened; as an exception, they rothney Cam., also studied by van der Vecht are brown-black, with or without narrow (1978, Bijdr. Dierk., 38/97-109); or of the yellow edge-bands, but never reddened‘. 17 subspecies of Polistes stigma (F.) of ‘Tegulae red or orange or light-yellow, Petersen (1987, Ent. Scand., 10:227-259); with transparent red superimposition ‘ or of the 9 ‘color forms’ of ‘Eumenes The new concept of subspecies of Car- lepeleterii Sauss.’ of Bequaert; or also 10 penter has hinged on not recognizing the ‘color varieties’ of Eumenes maxillosus validity of the subspecies, that are ‘artifi- (DG), also of the Bequaert (1918, Bull. Am. cial taxa’. This concept is taken from Car- Mus. nat. Hist., 39: 276-278 and 60: 279- penter in SPHECOS (1988, no. 17:12), who 281). It is opportune to remember how- approves the elimination of the subspecies ever how a lot of the ‘color varieties’ of proposed by the Archer ‘especially to the Bequaert are considered good species to- extent that subspecific taxa are eliminated’. day. And it is not impossible that some And reinforces this principle citing the re- subspecies of argillaceum will be revealed, search of McLean (really preceded by a lot to a more careful study, also valid species. of other varied authors, especially on Lepi- I can already say that the study of some doptera) who have ‘even shown experimen- copulatory apparatus of some has already tally that such ‘subspecies‘ in Polistes can put in evidence meaningful differences, be produced by manipulation of humidity!’. especially in the basal lobes of the aedeagus Really, the fact that varying some envi- and in the apodemes, as results from the ronmental factors induces variations of example of the figs.
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