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Comparisons of the Foveae Dorsales in Male and Female Ixodid Ticks Amblyomma Americanum, A. Maculatum, Dermacentor Andersoni

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Comparisons of the Foveae Dorsales in Male and Female Ixodid Ticks Amblyomma Americanum, A. Maculatum, Dermacentor Andersoni 0/2 J. Med. Entomol. Vol. 16, no. 3: 173-179 12 October 1979 @ 1979 by the Bishop Museum COMPARISONS OF THE FOVEAE DORSALES IN MALE AND FEMALE IXODID 0 TICKS AMBLYOMMA AMERICANUM, A. MACULATUM, DERMACENTOR ANDERSONI AND D. VARIABILIS (ACARI: IXODIDAE)1 By R. C. Axtell and A. LeFurgey2 Abstract. The presence of foveae dorsales and underlying microscopy. In the male tick the foveae dorsales foveal glands was documented in the adult males and compared to those in the females of Amblyomma americanum, A. maculatum, are even smaller and extremely difficult to locate Dermacentor andersoni and D. variabilis. Externally the male fo- even with scanning electron microscopy (SEM). veae dorsales were smaller and had fewer slits than the female Consequently, there have been considerable inad- glands, but internally there were no obvious gross differences equacies in published reports as to whether or not between the sexes in the foveal glands. the males possess foveae dorsales. Generally com- Several species of ixodid ticks have been shown prehensive publications on tick systematics and behaviorally to possess a sex pheromone produced morphology have mentioned or labeled foveae by the female and attractive to the males (Berger dorsales in simple diagrammatic drawings of the et al. 1971, Sonenshine et al. 1974). In the case of males as well as the females, often with the incor- 1 species, Amblyomma maculatum Koch, there is evi- rect assumption that they were sense organs (Ar- dence of production of a pheromone by the males, thur 1960, 1961, Cooley 1938, Dinnik & Zumpt since the females are attracted to the males (Glad- 1949, Douglas 1943, Hooker et al. 1912, Nuttall & ney 1971, Gladney & Drummond 1970a, b, Glad- Warburton 1911, Robinson 1926, Salmon & Stiles ney et al. 1974a, b). The female-produced sex 1901, Schulze 1942). Balashov (1968) mentioned pheromone in Amblyomma americanum (L.), Derma- foveae dorsales only in the females. With only the centor andersoni (Stiles) and D. variabilis (Say) is 2,6- light microscope available, it is questionable wheth- dichlorophenol (Berger 1972, Sonenshine et al. er these reports were based on observations of the 1976). The source of the sex pheromone in adult foveae dorsales or some other integumental light- female D. andersoni and D. variabilis is the foveal diffracting areas in the male ticks. Layton & So- glands beneath the foveae dorsales. These glands nenshine (1975) were unable to find the foveae were first described and their function elucidated dorsales in males of D. andersoni and D. variabilis. by Sonenshine and co-workers (Layton & Sonen- Subsequently, Sonenshine (pers. commun.) re- shine 1975, Sonenshine et al. 1977 , Vernick et al. ported finding them. In A. americanum and A. mac- 1978). ulatum the foveal glands have not been investigat- The foveae dorsales are paired, consist of a ed. However, Gladney (pers. commun.) observed group of external openings (slits) and are located what were likely foveae dorsales in the male A. on the dorsal integument usually near the midline maculatum. With the dramatic progress in research and in an area opposite approximately the mid- on the sex pheromone and behavior of these point between the 3rd and 4th coxae. However, species, there is a need for published illustrations due to their small size, the foveae dorsales are dif- and comparisons of the foveae dorsales in both L sexes and these are presented herein. ( ficult to find in the female tick with the aid of light MA TERIALS AND METHODS lPaper No. 5787 of the Journal Series of the North Carolina Agricultural Experiment Station. Raleigh, NC. For scanning electron microscopy of the exterior 2Professor and Research Associate, respectively, Dep. of structure of the foveae dorsales, unfed living Entomology, North Carolina State University, Raleigh, NC 27650, USA. adults of both sexes (10-26 specimens) of Amblyom- 174 J. Med. Entomol. Vol. 16, no. 3 TABLE1. Comparison of foveae dorsales in <3and 'i' of 4 Autoscan the dried sections were mounted on alu- species of ixodid ticks. minum stubs and coated with carbon-gold/pallad- SLIT LENGTH ium. Operating voltages were 5 to 15 kV. SEX DrAM (/.Lm)* No. SLITS* (,.Lm)* Amblyomma americanum RESULTS 'i' 36-78 (56) 10-30 (15) 2.4-7.1 (5.1) <3 17-23 (24) 4-11 (7) 1.9-4.6 (3.1) External structure (FIG. 1-8) Amblyomma maculatum In both females and males the foveae dorsales 'i' 45-84 (66) 25-28 (26) 3.8-6.0 (5.0) consist of paired depressions or pits, located at ap- <3 20-30 X 27-42 10-21 (16) 2.8-5.2 (3.8) (24 x 36) proximately % the length of the body from the Dermacentor andersoni capitulum to the posterior edge and centered from side to side on the dorsum. In the female the fo- 'i' 47-144 (98) 13-35 (26) 4.3-8.5 (5.4) <3 14-70 (52) 17-27 (22) 2.5-6.2 (4.2) veae dorsales are on the extensible alloscutal area Dermacentor variabilis posterior to the nonextensible scutum, approxi- 'i' 81-112 (97) 19-26 (22) 4.3-7.1 (5.4) mately over the genital pore, and at the level of <3 33-50 (41) 6-13 (12) 2.9-5.8 (4.6) the last pair of legs. In the male the foveae dorsales * Range (avg). occur on the nonextensible dorsal area, posterior to the pseudoscutum, and also are centered at the level of the last pair of legs. A single large seta ma americanum, A. maculatum, Dermacentor andersoni, usually occurs near each pit in both males and fe- and D. variabilis were coated with carbon-gold/pal- males. The foveae dorsales are circular in all ladium and examined with an ETEC Autoscan species examined except in the males of A. macu- scanning electron microscope at an operating volt- latum, in which they are distinctly oval. In males of age of 10 kV. the 4 species the diameter of each fovea is smaller For scanning electron microscopy of the interior than in females (TABLE 1), in proportion to the structure of the foveae dorsales, unfed living smaller overall body dimensions for males as com- adults of both sexes were fixed 12-24 h in Na-cac- pared to females. The nonextensible cuticle of the odylate-buffered 5% glutaraldehyde at 4°C. Small male is similar in texture to and contiguous with slits were cut in the edge of the cuticle of all spec- the dorsal pit. Therefore, the foveae dorsales are imens to allow penetration of the glutaraldehyde not as evident in males as in females, in which the into internal tissues. The whole ticks were then accordion folds of the extensible cuticle contrast washed 12-24 h in Na-cacodylate buffer at 4°C, sharply with the relatively smooth area of the pits dehydrated in a graded ethanol series of 15% to and emphasize the slight depression of the pits be- 100%, and sectioned through the foveae dorsales low cuticular level (e.g., see FIG. 9b, lOb). using a cryofracture technique adopted from Except in A. americanum males, the depressions Humphreys et al. (1974) and Lim (1971). A shal- are visible under the dissecting microscope at 50 X low aluminum dish containing a whole tick in in both sexes of all species. The average diameter 100% ethanol was placed on a liquid nitrogen- of a fovea in the A. americanum males examined cooled brass block, and positioned for viewing un- was 24 fLm,approximately 12 the size of a fovea in der a stereomicroscope. Liquid nitrogen was gently Dermacentor males and ~ to 12 the size of a fovea poured around the dish and block so that the tick in Dermacentor and Amblyomma females (TABLE 1). was frozen within the ethanol. The foveae dorsales The depressions in A. americanum males, while dif- remained visible through the frozen ethanol at ficult to locate initially, may be seen at 100 x. 50x in all species (except A. americanum males, With the scanning electron microscope, the 100 x), allowing horizontal or longitudinal frac- structures within the depressions of either Am- tures to be made through the foveae dorsales with blyomma or Dermacentor specimens are clearly visi- a nitrogen-cooled single-edge razor blade. The sec- ble. In males and females of the 4 species exam- tioned tick and surrounding ethanol were allowed ined, the depression is composed of irregularly to thaw and the tick sections were then transferred arranged slits, varying in both length and number to fresh 100% ethanol. The fractured segments from species to species. The orientation of the slits were critical point-dried in freon-13 after inter- is neither distinctly posterior-anterior nor distal- mediate replacement of the ethanol with freon-113 lateral; both directions and gradations from one to (Nemanic 1972). For observation in the ETEC the other direction, occur within a single fovea. 1979 Axtell & LeFurgey: Comparisons of foveae dorsales in ixodid ticks 175 - . ~ l.. ,.. .- . -.. I ~ ..~_. m ...~ OJ E. "- i!!II1 - .. '" - .-- I - n .... FIG. 1-4. L Adult 'i' Amblyomma americanum. la, dorsal view showing locations of foveae dorsales (arrows) (21 x). Ib, closer view of the pair of foveae dorsales (138 x). Ic, close-up of a fovea dorsales showing slit openings (892x). 2. Adult <3Amblyomma americanum. 2a, dorsal view showing locations of foveae dorsales (arrows) (l9x). 2b, closer view of the pair of foveae dorsales (44Ix). 2c, close-up of a fovea dorsales showing slit openings (l582x). 3. Adult 'i' Amblyomma maculatum. 3a, dorsal view showing locations of foveae dorsales (arrows) (14x). 3b, closer view of the foveae dorsales (l70x). 3c, close-up of a fovea dorsales showing slit openings (741 x).
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